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Full text of "Bulletin of the Natural History Museum"

Bulletin of _ 

The Natural History 



Bfit-RSH MU8&M 



PRIteifTBD 
QENERAl LIBRARY 



Botany Series 





VOLUME 23 NUMBER 2 25 NOVEMBER 1993 



The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum 
(Natural History)), instituted in 1949, is issued in four scientific series, Botany, 
Entomology, Geology (incorporating Mineralogy) and Zoology. 

The Botany Series is edited in the Museum's Department of Botany 
Keeper of Botany: Dr S. Blackmore 

Editor of Bulletin: Dr R. Huxley 

Assistant Editor: Mrs M.J. West 



Papers in the Bulletin are primarily the results of research carried out on the unique and ever- 
growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who 
make use of the Museum's resources. Many of the papers are works of reference that will remain 
indispensable for years to come. All papers submitted for publication are subjected to external peer 
review for acceptance. 

A volume contains about 160 pages, made up by two numbers, published in the Spring and Autumn. 
Subscriptions may be placed for one or more of the series on an annual basis. Individual numbers 
and back numbers can be purchased and a Bulletin catalogue, by series, is available. Orders and 
enquiries should be sent to: 

Intercept Ltd. 
P.O. Box 716 
Andover 
Hampshire SPIO lYG 

Telephone: (0264) 334748 
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Wo rW Lwr abbreviation: Bull. nat. Hist. Mus. Lond. (Bot.) 
© The Natural History Museum, 1993 



Botany Series 
ISSN 0968-0446 Vol. 23, No. 2, pp. 55-177 

The Natural History Museum 

Cromwell Road 

London SW7 5BD Issued 25 November 1993 

Typeset by Ann Buchan (Typesetters), Middlesex 
Printed in Great Britain at The Alden Press. Oxford 



Bull. nat. Hist. Mus. Lond. (Bot.) 23(2): 55-59 



Issued 25 November 1993 



New taxa of Gentiana (Gentianaceae) from 
Western China and the Himalayan region 



- 3 DEC 19 93 



TING-NUNG HO AND SHANG-WU LIU 

Northwest Plateau Institute of Biology, Academia Sinica, Xining, Qinghai, China 



PRISENTiO 

GENERAL LIBRAR 



Synopsis. Eleven new species* and two new varieties from Western China and the Himalayan region are described. 
They are G. leucantha H. Smith, G. chateriT. N. Ho, G. hicksii H. Smith, G. masoniiT. N. Ho, G. zekuensis T . N. 
Ho et S. W. Liu, G. laxiflora T. N. Ho, G. depressa var. stenophylla T. N. Ho, G. bryophylla H. Smith, G. micans 
var. latifolia T. N. Ho, G. glabriuscula H. Smith, G. winchuanensis T. N. Ho, G. shaanxiensis T. N. Ho and G. 
subuliformis S.W. Liu. The following three new names are also made: G. himalayaensis T. N. Ho, G. a/a/a T. N. Ho 
and G. taiwanica T. N. Ho. 

*Among them three species were diagnosed, described and typified and one species labelled by Harry Smith but 
were left unpublished at his death in 197L 



NEW TAXA 



I. Gentiana leucantha H. Smith, sp. nov. (Sect. Otophora 

Kusn.) 
Species Gentiana decoratae Diels et Gentiana infelici C. B. 
Clarke affinis, sed a prima tubo corollae lobis multo longiore 
(nee subaequilongo), a secunda flore duplo majore, ab ambo- 
bus flore albo (in sicco lutescenti) bene differt. 

Planta perennis, omnino glabra, e coUo radicem principalem 
carnosulam, fusiformem et caules floriferos numerosos 
edens. Caules floriferi perennes, assurgentes, breves, dense 
parvifoliati. Folia rosularia decunt, caulina sessilia, interno- 
diis multo longiora, ovata, 3.5-4 mm longa, 1.8-2 mm lata, 
apice acuta, submucronata, margine cartilagineo- 
microscabridula. Flores solitarii, terminales, erecti, sessiles; 
tubus calycis campanulatus, 5-6 mm longus, lobis ellipticis, 
3-3.5 mm longis, c. 2 mm latis, apice acuminatis, mucronatis; 
corolla alba (in sicco lutescens), calyce triple vel ultra lon- 
gior, tubo subcylindrico, 13-15 mm longo, lobis ovatis, 
4-5 mm longis, fere 4 mm latis, obtusis, plicae lobis later- 
aliter adnatae, c. 2.5 mm longae et 1.5 mm latae, bifidae, 
laciniatae vel fere integrae subtruncataeque; stamina tubo 
corollae c. 7 mm alte affixa, ex ore corollae exserta, filamen- 
tis filiformibus, 6-8 mm longis, antheris linearibus, vix 2 mm 
longis; ovarium ellipticum, 8-10 mm longum, apice in stylum 
c. 1 mm longum attenuatum, stipite c. 2 mm longo. Capsula 
inclusa, c. 11 mm longa; semina brunnea, oblonga, 1.1 X 
0.4 mm magna, tenuiter reiculata. 

China : S Tibet : Tsari , Karky n la , on open grassy hillside , 4420 m , 
24 June 1936, Ludlow & Sherriff 2209 (BM); ibid, on mossy open 
hillside, 4000-4270 m, 11 October 1938, Ludlow, Sheriff & 
Taylor 6592 (BM); Chayul chu, Kashong la, 4420 m, 14 July 
1936, Ludlow & Sherriff 2356 (BM); SE Tibet: Langgong, 28°45' 
N 94°E, on open grassy hillside, 4420 m, 5 June 1938, Ludlow, 
Sherriff & Taylor 3963 (BM, E); Kongbo, Kusha la near Paka, 
29°15' N 94°2' E, 4270-4570 m, 25 July 1938, Ludlow, Sherriff & 
Taylor 5934 (BM-holotype); ibid, on open grassy meadows, 4270 
m,' 27 July 1938, Ludlow, Sherriff & Taylor 5951 (BM, E); ibid. 
Deyang la, on stony grassy slopes, 4115 m, 8 August 1947, 



Ludlow, Sherriff & Elliot 14273 (BM, E). NE Bhutan: Me la, 
4420 m, 5 August 1933 Ludlow & Sherriff 415 (BM); ibid. 4270 
m, 26 August 1949, Ludlow, Sherriff & Hicks 21104 (BM, E, K); 
Ju la, Mangde chu, 19 July 1949, Ludlow, Sherriff & Hicks 16886 
(BM,E). 

2. Gentiana chateri T. N. Ho, sp. nov. (Sect. Otophora 

Kusn.) 
Species ob folia rosularia linearia et corollam albam usque 
flavescentem Gentiana damyonensi Marquand et Gentiana 
hicksii H. Smith similis, sed a prima corolla epunctata, lobis 
tubo brevioribus, foliis caulinis ovato-lanceolatis, a secunda 
corolla minore, 18-20 mm longa, ab ambobus floribus 3^ in 
cymam laxam dispositis, lobis calycis irregularibus, oblan- 
ceolatis et spathulatis, basi contractis recedit. 

Herba perennis, 3-4 cm alta, e collo radicem principalem, 
caules floriferos et rosulem basalem emittens. Radix carno- 
sula, verticalis, fusiformis, 4—6 mm diam. Caules floriferi 1^, 
ascendenti-erecti, glabri. Folia rosularia linearia vel lineari- 
oblonga, 8-22 mm longa et 2-4 mm lata, apice acuta vel 
obtusa, basi attenuata, nervis 1-3, subtus prominentibus, 
petiolis c. 2 mm longis; ilia caulina 2-3-juga, remota, ovato- 
lanceolata, 7-18 mm longa et 3-7 mm lata, apice acuta, basi 
breviter petiolata. Flores saepe 3^ in cymam laxam dispositi, 
raro solitarii; pedicelli 4-5 mm longi; calyx 7-9 mm longus, 
tubo tubuloso, lobis irregularibus, obovatis et lineari- 
spathulatis, 3-4 mm longis et 0.5-1.5 mm latis, apice acutis 
vel obtusis, basi contractis; corolla alba (flavescens in sicco), 
atrocaeruleo-striata, tubulosa, 18-20 mm longa, lobis tubo 
brevioribus, oblongis, 6-7 mm longis, acutis, plicae obliquae, 
auriculatae, lobis lateraliter adnatae; stamina tubo corollae 
inferioris inserta, filamentis subulatis, c. 10 mm longis, ad 
basin breviter ampliatis, antheris luteis, c. 1 mm longis; 
ovarium lineare, c. 15 mm longum, stylo c. 5 mm longo, 
stigmatibus linearibus, recurvatis. 

Nepal: Iswa Khola, 4000 m, 8 August 1971, L. W. Beer, C. R. 
Lancaster & D. Morris 9545 (BM), Kasuwa khola, 4000 m, 23 
August 1975, L. W. Beer 25363 (BM-holotype). 

We take particular pleasure in naming this species after Mr 



©The Natural History Museum, 1993 



56 

Arthur O. Chater, former curator of the flowering plant 
herbarium, The Natural History Museum, who worked for 
some years on the Nepalese Flora. 

3. Gentiana hicksii H. Smith, sp. nov. (Sect. Otophora 
Kusn.) 

Species Gentiana damyonensi Marquand affinis, sed corolla 
majore, 25-35 mm longa, tubo lobis longiore (nee breviore) 
differt. 

Herba perennis, c. 8 cm alta, e collo radicem principalem, 
caules floriferos et rosulam basalem emittens. Radix carno- 
sula, verticalis, fusiformis, 3-5 mm diam. Caules floriferi 1-3, 
ascendenti-erecti, glabri. Folia rosularia linearia vel 
lanceolato-linearia, 3-4 mm longa et 2-3.5 mm lata; ilia 
caulina 3-4-juga, internodiis breviora vel interdum sub- 
aequilonga, lanceolato-linearia, 0.7-1.2 cm longa, acuta, ses- 
silia. Flores saepe solitarii, terminales; calyx 7-9 mm longus, 
tubo obconico, lobis subregularibus, linearibus; corolla alba, 
coeruleo-striata, cylindrico-obconica, 25-35 mm longa, tubo 
c. 19 mm longo, lobis ovatis, 8-9 mm longis et 5 mm latis, 
obtusis, plicae dentiformes, latere lobi adnata; stamina tubo 
c. 6 mm alte affixa, filamentis 13-15 mm longis, antheris 
angustis, vix 2 mm longis; ovarium lanceolatum, c. 15 mm 
longum, apice in stylum c. 4 mm longum attenuatum, stipite 
2-3 mm longo. Capsula et semina non visa. 

Bhutan: Pang la, on open wet slopes, 4000 m, 21 September 
1949, Ludlow, Shernff& Hicks 21456 (BM - holotype; UPS - 
isotype). 

4. Gentiana masonii T. N. Ho, sp. nov. (Sect. Otophora 
Kusn.) Fig. 1. 

Species Gentiana otophorae Franchet ex Forbes & Hemsley 
affinis, sed corolla omnino coerulea, epunctata bene differt. 

Herba perennis, omnino glabra, 14-18 cm alta, e collo radi- 
cem principalem, caules floriferos et rosulam basalem emit- 
tens. Radix carnosula, cylindrica, 4-7 mm diam. Caules 
floriferi 1-3, ascendenti erecti. Folia rosularia oblongo- 
spathulata vel oblonga, 3-7.5 cm longa et 0.9-1.9 cm lata, 
apice acuta vel obtusa, basic in petiolos 2.5-6 cm longos 
attenuata, nervis 3-5, subtus prominentibus, ilia caulina 
2-3-juga, remota, oblonga vel ovato-elliptica, 1.4—2.6 cm 
longa et 0.7-1.2 cm lata, apice obtusa vel acuta, basi in 
petiolos 3-7 mm longos attenuata. Flores 3^ in cymam 
laxam dispositi; calyx 6-8 mm longus, lobis irregularibus, 
spathulatis, 1.5-2.5 mm longis apice rotundatis vel acutis, 
basi contractis; corolla caerulea, epunctata, cylindrica, 
16-20 mm longa, lobis tubo longioribus, oblongis, 13-16 mm 
longis et c. 5 mm latis, apice acutis, plicae minimae, c. 1 mm 
longae, auriculatae, lobis lateraliter adnatae; stamina medio 
corollae inserta, filamentis subulatis, c. 10 mm longis, anth- 
eris caeruleis, c. 1.5 mm longis; ovarium cylindricum, 
11-13 mm longum, breviter stipitatum, stylo breve, stigmate 
breve. 

Upper Burma: N'Maikha-Salwin divide, 26°28' N 98°48' E, 
September 1924, Forrest 24944 (K); ibid. 26°50' N 98°48' E, 
on alpine meadows, 4270 m, September 1925, Forrest 27222 
(BM - holotype; K - isotype). 

This new species is dedicated to my friend, Prof. Howard S. 
Mason, a member of American Rock Garden Society. He has 
been devoting himself to cultivated Gentians of different 
Gentian seedUngs. 



T -N. HO AND S -W. LIU 

5. Gentiana zekuensis T. N. Ho & S. W. Liu, sp. nov. (Sect. 
Cruciata Gaudin) Fig. 2. 

Species habitu Gentiana officinali H. Smith valde similis, sed 
tubo calycis obtuso (nee truncate), lobis filiformibus, usque 
ad 5 mm longis (nee dentiformibus, 1-1.5 mm longis), corolla 
flavida, caeruleo-punctata, versus limbum caerulea, raro 
corolla omnino caerulea (nee omnino flavida) differt. 

Herba perennis, 15-40 cm alta. Radices subcarnosae, pau- 
cae, in radicem verticalem, cylindricam conniventes. Caules 
floriferi 3-10, ascendentes, caudice reliquis petiolorum bru- 
neis, fibrosis obtecto. Folia rosularia anguste elliptica vel 
lineari-lanceolata, 8-22.5 cm longa et 0.5-2.5 cm lata, apice 
acuta, basi attenuata, utrinque glabra, nervis 3-5, subtus 
prominentibus; ilia caulina 2-3-juga, elliptica vel lanceolata, 
1.8-5.5 cm longa et 0.6-1.2 cm lata, apice acuta, basi in 
vaginam 0.8-3 cm longam connata, eis summis bractescenti- 
bus, lanceolatis usque lineari-lanceolatis, 1.2-5 cm longis et 
0.5-1 cm latis, acuminatis. Flores sessiles, numerosi, termina- 
les axillaresque; calyx fissus fere ad basin, 6-13 mm longus, 
tubo spathaceo, apice obtuso, lobis filiformibus, usque 5 mm 
longis; corolla flavida, caeruleo-punctata, versus limbum 
caerulea, raro corollla omnino caerulea, tubulosa, lS-20 mm 
longa, lobis ovatis, 2.5-3 mm longis, acutis, plicae obliquo- 
triangulatae, c. 1 mm longae, acuminatae; stamina tubo 
corollae inferioris inserta, filamentis linearibus, 7-8 mm 
longis, antheris flavidis, Hnearibus, c. 1.5 mm longis; ovarium 
lanceolatum, 11-13 mm longum, stylo cum stigmatibus c. 
2 mm longo. 

The corollas of G. zekuensis are yellowish, bluish towards the 
limb, seldom whole corolla bluish. G. zekuensis is somewhat 
intermediate between G. macrophylla and G. officinalis but 
closer to the latter. It differs from both mainly by the distinct 
and filiform calyx lobes. 

China: Qinghai. Zeku, among shrubs, 3600 m, 24 July 1967, 
Y. C. Young 1822 (HNWP); ibid, on grassy hillside, 3200 m, 
27 August 1970, L. H. Zhou & L. N. Sun 1958 (HNWP - 
holotype); ibid, among shrubs, 3300 m, 14 August 1975, Z. 
B. Wang 349 (HNWP); Tongren among shrubs, 3400 m, 26 
July 1970, 5. W. Liu 1474 (HNWP). 

6. Gentiana laxiflora T. N. Ho, sp. nov. (Sect. Frigida Kusn.) 
Species G. trichotomae Kusn. et G. erecto-sepalo T. N. Ho 
affinis, ab hac corolla 22-25 mm longa, lobis calycis angustis, 
linearibus et subulatis, sinubus inter lobos calycis rotundatis, 
ab ilia corolla caerulea, versus basin flavescente differt. 

Herba perennis, 10-14 cm alta. Rhizoma breve, horizontale 
vel verticale, radices adventitias, caules floriferos et rosulam 
basalem emittens. Caules 1-3, erecti, subglabri, basi vaginis 
veteribus membranaceis obtecti. FoHa plerumque basilaria, 
petiolata, laminae lineari-oblongae, 2.5-8 cm longae et 
3-6 mm latae, apice acutae vel obtusae, basi attenuatae; ilia 
caulina 1-2 juga, minora, sessilia. Flores 5-7, terminales et 
axillares, in florescentiam laxam dispositi; pedicelli graciles, 
5-21 mm longi; calyx tubulosus, 8-11 mm longus, lobis 
angustis, linearibus et subulatis, 3-6 mm longis, sinubus inter 
lobos calycis rotundatis; corolla caerulea, versus basin flave- 
scens, infundibuliformis, 2-25 mm longa, lobis ovatis, 
1.5-3 mm longis, acutis, plicae truncatae, lobis breviores; 
stamina tubo corollae inferioris inserta, filamenti subulatis, c. 
10 mm longis, antheris luteis, c. 2 mm longi. Capsula breviter 



NEW TAX A OF GENTIAN A 



57 




cm 



cm 



cm 



Fig, 1 Gentiana masonii T. N. Ho: a, habit ; b, cauline leaves ; c, calyx opened out; d, corolla opened showing stamens. 



stipitata, ex corolla paullo exserta. 

China. SE Tibet. Nam la, 29°59' N 94°17' E, 4115 m, 30 
August 1938, Ludlow, Sherriff & Taylor 6944 (BM), Kongbo, 
30°N 94°20' E, 4115 m, 20 July 1947, Ludlow, Sherriff & 
Elliot 15492 (BM - holotype; E - isotype). 

7. Gentiana himalayaensis T. N. Ho, stat. et nom. nov. 

Gentiana nubigena var. parviflora C. B. Clarke in Hook, f., 
Fl. Brit. India 4: 117 (1883). Type: Sikkim, Kinchinjhow, 
4880-5200 m, /. D. Hooker (K - holotype!; BM - iso- 
type!) 

Gentiana algida var. parviflora (C. B. Clarke) Kusn. in Acta 
Horti Petrop. 15: 266 (1898). 

Distribution. C. & E. Himalayan region. China (S. &. SE 
Tibet), Nepal, Sikkim, Bhutan. 

8. Gentiana depressa D. Don var. stenophylla T. N. Ho, var. 

nov. (Sect. Isomeria Kusn.) 
A var. typo recedit foliis angustis, lineari-oblongis vel lan- 
ceolatis, 1-3.5 cm longis et 2.5-4(-8) mm latis, acuminatis. 

Leaves linear-oblong or lanceolate; calyx-lobes narrowly tri- 
angular or linear-elliptical. 

Nepal: East of Chalike Pahar, 4000 m, Stainton, Sykes & 
Williams 4593 (BM); Moktintan, on grassy slopes near Pass, 



4000 m, 1 October 1954, Stainton, Sykes & Williams 8060 
(BM); Phagune Dhuri, 4000 m, 13 October 1954, Stainton, 
Sykes & Williams 9007 (BM); Kali Gandaki, Tasang/ESE of 
Tukche, 3840 m, 4 November 1976, G. Miehe 194 (BM); S. 
side Deorali ridge, 2900 m, 11 November 1979, A. D. 
Schilling 2445 (BM - holotype). 

9. Gentiana alata T. N. Ho, nom. nov. 

Gentiana kusnezowii Franchet in Bull. Soc. bot. Fr. 43: 492 
(Nov. 1896), non Gilg May 1896 (in Engler, Bot. Jahrb. 
22: 325). Type: China, Yunnan, Yunnansen (Kunming), 
Delavay s.n. (P - holotype!). 

Distribution. China (C. Yunnan). 

10. Gentiana bryophylla H. Smith, sp. nov. (Sect. Chondro- 
phylla Bunge) 

Species habitu, forma folii et colore magnitudineque floris 
Gentiana burmensi Marquand persimilis, sed plicis corollae 
margine crenulatis, efimbriatis valde differt. 

Herba perennis, c. 5 cm alta, stolone breve praedita. Caulis 
ascendens vel erectus, uniflorus, subglaber. Folia basilaria 
minuta, 2-3 mm longa, 1-1.5 mm lata; ilia caulina 6-7-juga, 
inferiora minora, superiora ad 5 mm longa et 2 mm lata, 
lanceolata, recurvato-aristata, margine cartilaginea, subgla- 
braque, versus basin gradatim dense ciliolata. Tubus calycis 



58 



T.-N. HO AND S.-W. LIU 




Fig. 2 Gentiana zekuensis S. W. Liu: a, habit ; b, flower; c, calyx opened out; d, corolla opened inside with stamens and gynoecium. 



6-7 mm longus, lobis 3.5 mm longis, acicularibus, subarista- 
tis; corolla purpurea, c. 17 mm longa, tubo c. 15 mm longo, 
lobis paullo obliquis, rotundato-triangularibus, acutis, apicu- 
latis, 2.5 mm longis et latis, plicae lobis paullo breviores, 
subrotundatae, margine crenulatae; stamina tubo corollae 
6 mm alte adnata, filamentis filiformibus, 4.5 mm longis, 
antheris c. 1 mm longis; ovarium stipitatum, obovato- 
ellipsoideum, c. 5 mm longum, stylo fere 2 mm longo. Cap- 
sula amplanato-obovata, c. 5 mm longa, alata; semina 
brunnea, subangulariter ellipsoidea, 0.7 X 0.3 mm magna, 
reticulata. 

Burma: Nam Tamai valley, 28° N 97°45' E. , on steep slopes in 
the mossy carpet underneath Arundinaria and Rhododen- 
dron, 3000 m, 8 September 1937, Kingdon Ward 13184 (BM- 
holotype); ibid. 9 Septemper 1937, Kingdon Ward 13208 
(BM). 

11. Gentiana micans C. B. Clarke var. latifolia T. N. Ho, 

var. nov. (Sect. Chondrophylla Bunge) 
Gentiana metaensis H. Smith in herb. 

A var. typo recedit foliis caulinis latioribus, ovato-lanceolatis 
usque lineari-lanceolatis, patentibus (nee acicularibus, caule 
fere adpressis), lobis calycis brevioribus, latioribus, breviter 
acutis (nee longe acuminatis), plicis bifidis (nee integris 
minute laciniatis). 



Bhutan: Liaru Thang, Ringchen chu, 3750 m, 8 June 1937, 
Ludlow & Sheiriff 3531 (BM); Thampe la, on bare hillside, 
4570 m, 14 August 1949, Ludlow, Sherriff & Hicks 17134 
(BM), Shingbe (Me la), 4115 m, 8 September 1949, Ludlow, 
Sherriff & Hicks 21166 (BM - holotype; UPS - isotype). . 

12. Gentiana glabriuscula H. Smith ex T. N. Ho, sp. nov. 

(Sect. Chondrophylla Bunge). 
Species Gentiana pedicellatae (G. Don) Griseb. similis, sed 
planta omnino glabra, foliis basilaribus majoribus, involucri- 
formibus, orbicularibus vel late oblongis, illis caulinis minori- 
bus, ovatis valde differt. 

Planta annua vel biennis, 2-5 cm alta, omnino glabra. Caulis 
gracilis, ascendens, e basi ramosus quasi caespitosus. Folia 
basilaria majora, involucriformia, orbicularia vel late oblonga, 
9-26 mm longa, 7-15 mm lata, apice rotundata, mucronata, 
basi in petiolos 3-5 mm longos abrupte contracta, nervis 1-3, 
utrinque prominentibus; ilia caulina minora, ovata, 3-5 mm 
longa et 2-3.5 mm lata, apice acuta mucronataque, basi leviter 
attenuata, subsessilia. Flores sessiles, solitarii ad apices ramu- 
lorum siti; calyx campanulatus, 3.5-5 mm longis, lobis ovatis, 
2-3 mm longis et 1.5-2 mm latis, apice acutis mucronatisque, 
basi contractis, recurvatis; corolla caeruleo-grisea, extra atro- 
caerulea, fauce nigro-punctata, campanulata, 5-7 mm longa, 
calycem paullo superans, lobis ovatis, c. 2 mm longis, apice 



NEW TAXA OF GENTIAN A 



59 



acutis mucronatisque, plicae late ovatae, c. 1 mm longae, lobis 
breviores, acutae, integrae vel bilobatae; stamina medio corol- 
lae inserta, filamentis c. 1 mm longis; ovarium obovato- 
oblongum cum stipite c. 4 mm longum, stylo breviore, 
stigmatibus oblongis. 

Bhutan: Trashi Yanasi chu, on mossy rocks and banks in 
mixed forest, 2590 m, 5 September 1949, Ludlow, Sherriff & 
Hicks 20608 (BM - holotype). Sikkim: Gangtok-Karponang 
road, F. H. Lister 9 (K). China: S Tibet: Monyul, 2 April 
1936, Ludlow & Sherriff 1264 (BM). 

13. Gentiana taiwanica T. N. Ho, stat. et nom. nov. 

Gentiana scabrida var. angusta Masam. in Trans, nat. Hist. 
Soc. Formos. 29: 64 (1939). Type: China, Taiwan, Nan- 
hutashan, Taihoku-syu, 15 July 1931, Masamune & K. 
Mori s.n. (herb. Taihoku Imperial Univ. Taiwan - holo- 
type, not seen). 

Gentiana angusta (Masam.) Liu et Kuo in Bull. Exp. Forest 
Natn. Taiwan Univ. 114: 176, pi. 4 (1974), non M. E. 
Jones 1908 (in Contrib. Western. Bot. 12: 52). 

Distribution. China (Taiwan). 

14. Gentiana winchuanensis T. N. Ho, sp. nov. (Sect. Chon- 
drophylla Bunge). 

Species Gentiana piasezkii Maxim, capsula anguste oblonga, 
calyce carinato-alato affinis, sed foliis ovatis, lobis calycis 
triangularibus, multo brevioribus bene recedit. 

Herba annua, 8-10 cm alta. Caulis ascendens, ramosus. Folia 
basilaria ignoti; ilia caulina remota, internodiis breviora, 
ovata, 4-6 mm longa et 2-4 mm lata, apice acuta, basi in 
petiolos 2-3 mm longos abrupte contracta, margine carti- 
laginea ciliolataque, utrinque glabra. Flores subsessiles, soli- 
tarii ad apices ramulorum siti, calyx tubuloso- 
infundibuliformis, 13-17 mm longus, lobis triangularibus, 
1.5-2 mm longis, basi c. 1 mm latis, acuminatis, margine 
cartilagineis, sinubus rotundatis; corolla caeruleo-purpurea, 
fauce flavida et nigro punctata, anguste infundibuliformis, 
25-30 mm longa, lobis ovatis, 6-7 mm longis, apice acutis 
mucronatisque, pHcae ovatae, lobis paullo breviores, denticu- 
latae, stamina medio corollae inserta, filamentis filiformibus, 
c. 5 mm longis, antheris c. 1 mm longis. Capsula inclusa, 
fusiformis, c. 10 mm longa, alata, stipite c. 5 mm longo; 
semina brunnea, oblonga, c. 1 mm longa, tenuiter reticulata. 

China: N Sichuan. Winchuan, 2400 m, in forest, 30 July 1975, 
Sichuan Veget. Exped. 8522 (HNWP- holotype). 

15. Gentiana shaanxiensis T. N. Ho., sp. nov. (Sect. Chon- 
drophylla Bunge). 

Species habitu Gentiana piasezkii Maxim, similis, sed foliis 
caulinis linearibis lobis calycis linearibus, usque ad fauce 
corollae, lobis corollae acuminato-caudatis differt. 

Herba annua, 7-12 cm alta. Caulis purpureus, ascendens, 
dense papillosus, e basi ramosissimus. Folia apice acuta vel 
acuminata, margine dense papillosa, utrinque glabra, costis 
subtus prominentibus; ilia basilaria majora, anguste oblonga, 
25-42 mm longa et 3.5-8 mm lata, sessilia vel subsessiHa; ilia 
caulina linearia, 9-22 mm longa et 1.5-3 mm lata, sessilia. 
Flores solitarii ad apices ramulorum siti; pedicelli 2-10 mm 
longi, dense purpureo-papillosi, calyx tubuloso infundibuli- 
formis, 20-27 mm longus, usque ad faucem corollae, tubo 



carinato alato, lobis linearibus, 6-8 mm longis; corolla pur- 
purea, hypocrateriformis, 25-35 mm longa, lobis anguste 
lanceolatis, 5-9 mm longis, basi 2-3 mm latis, apice 
acuminato-caudatis, plicae ovatae, 4—5 mm longae et c. 3 mm 
latae, acutae, denticulatae; stamina medio corollae inserta, 
filamentis filiformibus, 50-55 mm longis, antheris linearibus, 
c. 1 mm longis; ovarium ellipticum, 7-8 mm longum, stylo 
cum stigmatibus 4-5 mm longo. Capsula e corolla exserta, 
spathulato-oblonga, 9-11 mm longa, alata, stipite corollam 
aequante vel superante; semina brunnea, oblonga, 1-1.2 mm 
longa, tenuiter reticulata. 

China: Shaanxi: Ningqiang, 11 July 1957, Northwest Univ. 
Biol. Exped. 57 (HNWP - holotype). 

16. Gentiana subuliformis S. W. Liu, sp. nov. (Sect. Frigida 

Kusn.) 
Species Gentiana wilsonii Marquand similis, sed foliis basi- 
laribus latis, anguste ellipticis vel oblongis, corolla multo 
minore, 23-25 mm longa differt. 

Herba perennis, 12-20 cm alta. Rhizoma breve, horizontale 
vel verticale, radices adventitias carnosulas, caules floriferos 
et rosulam basalem emittens. Caules floriferi erecti, basi c. 
2 mm diam., vaginis veteribus membranaceis obtecti. Folia 
plerumque basilaria, petiolata, petiolis 0.5-2 cm longis, lami- 
nis anguste ellipticis vel oblongis, raro oblongo-spathulatis, 
1.5-3.5 cm longis et 0.5-0.9 cm latis, obtusis vel subrotunda- 
tis, basi attenuatis, nervis 3, subtus prominentibus; ilia 
caulina 1-2-juga, eis basilaribus similaria, 2-3.5 cm longa et 
ad 1.2 cm lata, sessilia vel breviter petiolata. Flores 
numerosi, terminales, aggregati in inflorescentiam subcapi- 
tatam; calyx tubulosus, c. 10 mm longus, saepe fissus, lobis 
recurvatis vel patentibus, subulatis, 1-2 mm longis; corolla 
atrocaerulea, epunctata, tubulosa, 23-25 mm longa, lobis 
ovatis, 1.5-2 mm longis, obtusis, plicae lobis breviores, trun- 
catae, integrae vel denticulatae; stamina tubo corollae inferi- 
oris inserta, filamentis subulatis, c. 9 mm longis, antheris 
luteis, linearibus, c. 1.5 mm longis; ovarium stipitatum, lin- 
eare, c. 20 mm longum, stylo breve, stigmatibus linearibus. 

China: SE Tibet: Bai ma, on alpine meadows, 4800 m, 10 
August 1973, Northwest Inst. Biol. Tibet Exped. 1367 
(HNWP - holotype); Bamda-Zogong, 4000 m, 5 September 
1977, P. C. Kuo & W. Y. Wang 23411 (HNWP). 



Acknowledgements. We are particularly indebted to the directors 
and staff of the herbaria of The Natural History Museum, the Royal 
Botanical Gardens, Kew, and the Royal Botanical Garden at Edin- 
burgh for permission to consult their collections, for facilities pro- 
vided and for the loan of specimens. We must also thank Prof. Tang 
Yan-cheng, Mr A O Chater and Dr Mike Gilbert for useful discus- 
sions in the preparation of this publication. 



REFERENCES 



Ho, T.-N. & Liu, S.-W. & Wu, C.J. 1988, Gentianaceae. Flora Reipublicae 

Popularis Sinicae 62. Beijing. 
Kusnezow, N. I. 1894. Subgenus Eugentiana of genus Gentiana Tourn. Trudy 

imp. S.-Peterb. Obshch. Estest. 24(2): 1-531. [Latin translation in Trudy 

imp. S. -Peterb. bot. Sada 15: 1-507 (1896-1904)]. 
Naithani, H. B. 1990. Gentiana. In Flowering plants of India, Nepal and 

Bhutan: 291-295. 



Bull. nat. Hist. Mus. Lond. (Bot.) 23(2): 61-65 



Issued 25 November 1993 



New combinations, names and taxonomic 
notes on Gentianella (Gentianaceae) from 
South America and New Zealand 

TING-NUNG HO AND SHANG-WU LIU 

Northwest Plateau Institute of Biology, Academia Sinica, Xining, Qinghai, China 

Synopsis. Gentianella is a large genus of about 200 species. It was segregated from the genus Gentiana in 1794. 
Since then the study histories of the two genera have been interwoven. During study on Gentiana and Gentianella it 
was realized that 104 new combinations and two names needed to be made from South America and New Zealand. 



INTRODUCTION 



The genus Gentianella was segregated from the genus Genti- 
ana L. by Moench in 1794. Since then the study histories of 
the two genera have been closely interwoven. The majority of 
species of Gentianella were originally described within Genti- 
ana. Since the last half century, Fabris (1953-1981), Gillet 
(1957), Holub (1967-1983) and Pringle (1981-1987) etc. have 
studied Gentianella, and made a number of nomenclatural 
transfers, but these studies are insufficient for the whole 
genus and there are still too many species under Gentiana. 
During study on Gentiana and Gentianella in Eurasian her- 
baria, it was realized that 104 new combinations and two 
names of Gentianella needed to be made from South America 
and New Zealand. 

Gentianella Moench is a large genus of about 200 species. It 
has an almost global distribution chiefly centred on South 
America and New Zealand. 



NEW COMBINATIONS AND TAXONOMIC 
NOTES 

Gentianella achalensis (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana achalensis Hieron., Bot. Acad. Nac. Cordova, 4: 373 

(1881), nom. nud; Gilg, Bot. Jahrb., 11: 322 (1896). Type: 

Argentina, Hieronymus 526 (K!). 

Gentianella albido-caerulea (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana albido-caerulea Gilg, Bot. Jahrb. ,11: 323 (1896). Syntypes: 
Bolivia, M. Bang 1132 (n.v.), O. Kuntze (n.v.). 

Gentianella andreae-mathewsii (Briquet) T. N. Ho & S. W. Liu, 
comb. nov. 

Gentiana andreae-mathewsii Briquet, Candollea, 4: 326 (1931), based 
on Gentiana mathewsii Gilg, Bot. Jahrb., 54 (Beibl. 118): 64 
(1917), non Petrie (1911) {Trans. Proc. N. Zeal. Inst., 44: 183). 
Type: Peru, Mathews (n.v.). 

Gentianella antarctica (Kirk) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana concinna var. robusta Hook, f., Fl. Antarct., 1: 53 (1844). 
Gentiana antarctica Kirk, Trans. Proc. N. Zeal. Inst. 27: 339, 1894 
(1895). Type: New Zealand, T. Kirk, W. 4729 (n.v.). 

Gentianella antipoda (Kirk) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana antipoda Kirk, Trans. Proc. N. Zeal. Inst., 23: 440. 1890 



(1891), nom. nud. et 27: 340. 1894 (1895). Type: New Zealand, T. 
Kirk (K! - isotype). 

Gentianella armerioides (Griseb.) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana armerioides Griseb. in Lechler, Berb. Am. Austr., 58 
(1857). Type: Peru, Lechler 2000a (P! - isotype). 

Gentianella astonii (Petrie) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana astonii Petrie, Trans. Proc. N. Zeal. Inst., 48: 187. 1915 

(1916). Type: New Zealand, Valley of Ure river, B. C. Aston 

(WELT! - holotype). 

Gentianella atroviolacea (Gilg) Fabris ex T. N. Ho & S. W. Liu, 
comb. nov. 

Gentiana atroviolacea Gilg, Bot. Jahrb., 54(Beibl. 118): 53 (1917). 

Type: Columbia, Kalbreyer 1200 (K! - isotype). 
Gentiana solidagoides Reim., Bot. Jahrb., 62: 329 (1929). Type: 

Columbia, (K! no. 10574), syn. nov. 

Gentianella bangii (Gilg) Fabris ex T. N. Ho & S. W. Liu, comb. nov. 
Gentiana bangii Gilg, Bot. Jahrb., 11: 324 (May 1896), nee Rusby 

(end 1896) {Mem. Torr. Bot. Club , 6: 79), Type: Bolivia, M. Bang 

1153 (K!- isotype). 

Gentianella bellatula (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana bellatula Gilg, Bot. Jahrb., 50(Beibl. HI): 49 (1913). 
Syntypes: Bolivia, Hauthal 201, 218 (all n.v.). 

Gentianella bellidifolia (Hook, f.) Holub var. divisa (Kirk) T. N. Ho 

& S. W. Liu, comb. nov. 
Gentiana bellidifolia var. divisa Kirk, Trans. Proc. N. Zeal. Inst., 27: 

337. 1894 (1895). Type: New Zealand, Ashburton Mountains, T 

H. Potts, W. 4714 (n.v.). 
Gentiana divisa (Kirk) Cheeseman., Man. N. Zeal FL, 453 (1906). 

Gentianella bockii (Gilg) T. N. Ho & S. W. Liu, comb, nov, 
Gentiana bockii Gilg, Bot. Jahrb., 54(Beibl. 118): 34 (1917). Syn- 
types: Bolivia, Bock in herb. Herzog 2480e (n.v.), Buchtien 1481 
(n.v.). 

Gentianella bridgesii (Gilg) Fabris ex T. N. Ho & S. W. Liu, comb, 
nov. 

Gentiana bridgesii Gilg, Bot. Jahrb., 11: 316 (1896). Syntypes: 
Bolivia, Bridges a. 1850 (n.v.), O. Kuntze (n.v.). 

Gentianella briquetiana (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana briquetiana Gilg, Bot. Jahrb., 54 (Beibl. 118): 32 (1917). 

Type: Bolivia, Larecaja, Mandon 361 (P! - holotype, BM! K! - 

isotypes). 

Gentianella bromifolia (Griseb.) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana bromifolia Griseb., Glett. Abh., 19: 208 (1874). Syntypes: 
Argentia, Tucuman, Lorrentz 775 (K!), 303 (n.v.). 

Gentianella brunneo-tincta (Gilg) T. N. Ho & S. W. Liu, comb. nov. 



)The Natural History Museum, 1993 



62 



T.-N. HO ANDS.-W. LIU 



Gentiana brunneo-lincta Gilg, Fedde, Rep. Nov. Sp., 2: 37 (1906). 
Type: Peru, Weberbauei 3092 (n.v.). 

Gentianella buchtienii (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana buchtienii Gilg, Bot. Jahrb., 54(Beibl. 118): 66 (1917). 

Type: Bolivia, Buchtien 48 (K! - lectotype designated here, BM! - 

isolectotype). 

Gentianella calcarea (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana calcarea Gilg, Fedde, Rep. Nov. Sp., 2: 42 (1906). Type: 
Peru, Weberbauer 2539 (n.v.). 

Gentianella centamalensis (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana centamalensis Gilg, Bot. Jahrb.,12: 334 (1896). Type: Peru, 
Stuebel 41 (n.v.). 

Gentianella cerina (Hook, f.) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana cerina Hook, f., Fl. Antarct., 1: 54, t. 36 (1844). Type: New 
Zealand, Ins. Auckland, J. D. Hooker (K! - holotype). 

Gentianella chathamica (Cheesman) T. N. Ho & S. W. Liu, comb, 
nov. 

Gentiana chathamica Cheeseman, Man. N. Zeal. FL, 449 (1906). 
Type: New Zealand, F. A. D. Cox (BM! K! - isotypes). 

Gentianella chrysantha (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana chrysantha Gilg, Bot. Jahrb., 54(Beibl. 118): 81 (1917). 
Type: Bolivia, Herzog 2044 (n.v.). 

Gentianella chrysosphaera (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana chrysosphaera Gilg, Bot. Jahrb., S4{BQ\b\. 118): 37 (1917). 
Type: Peru, Weberbauer 6521 (n.v.). 

Gentianella chrysotaenia (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana chrysotaenia Gilg, Bot. Jahrb., 54(Beibl. 118): 39 (1917). 
Syntypes: Peru, Weberbauer 652S, 6914 (all n.v.). 

Gentianella claytonioides (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana claytonioides Gilg, Bot. Jahrb., 22: 318 (1896). Type: 
Argentina, Rioja, Hieronymous & Niederlein (n.v.). 

Gentianella coccinea (G. Don) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana coccinea G. Don, Gen. Syst., 4: 196 (1838). Type: Peru, 
Pavon (BM! - isotype). 

Gentianella concinna (Hook, f.) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana concinna Hook, f., Fl. Antarct.,53, t. 35 (1844). Type: New 
Zealand, Ins. Auckland, J. D. Hooker (K! - holotype). 

Gentianella coquimbensis (Briquet) T. N. Ho & S. W. Liu, comb, 
nov. 

Gentiana coquimbensis Briquet, Candollea, 4: 328 (1931). Type: 
Chile, Coquimbo, M. Gay (P! - isotype) 

Gentianella corallina (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana corallina Gilg, Fedde, Rep. Nov. Sp., 2: 48 (1906). Type: 
Peru, Weberbauer 4288 (n.v.). 

Gentianella crassiuscula T. N. Ho & S. W. Liu, nom. nov. 
Gentiana crassicaulis Gilg, Bot. Jahrb., 54(Beibl. 118): 60 (1917), 

non Duthie ex Burkill (1906) (7. As. Soc. Beng. n. s. 2: 316). Type: 

Peru, Lobb (n.v.). 

Gentianella crassulifolia Fabris, Bol. Soc. Argent. Bot., 8: 25 (1959). 
Gentiana crassulifolia Griseb., Gen. Sp. Gent., 227 (1838). Type: 

Columbia, Jameson 455 (K!). 
G. crassulifolia is a subshrub, characterised by crowded, imbricate, 
rigid and subcoriaceous leaves, very short pedicels and flowers 
crowded in a head at the top of the stem. Its calyx-lobes are as long as 
to slightly longer than the tube. Leaf shape shows some variation. 
They are oblong, elliptic, lanceolate to ovate. Two varieties can be 
recognised. 

var. crassulifolia 

Gentiana selaginifolia Griseb., Linnaea, 11: 42 (1849). Type: Colum- 
bia, Hartweg 1252 (P! - isotype). 

Gentiana engleri Gilg, Bot. Jahrb., IT. 314 (1896). Type: S. Colum- 
bia, Lehman 666 (BM!, K!), syn. nov. 

Gentiana dacrydioides Gilg, Bot. Jahrb., 11: 312 (1896). Type: S. 
Columbia, Lehman 2682 (BM! - isotype), syn. nov. 



Gentianella selaginifolia (Griseb.) Fabris, Bol. Soc. Argent. Bot., 8: 

25 (1959). 
Gentianella dacrydioides (Gilg) Weaver & Rudenberg, J. Arnold. 

Arbor., 56(2): 215 (1975). Calyx-lobes lanceolate; corolla c. 

15 mm. 

var. hypericoides (Gilg.) T. N. Ho, stat. et comb. nov. 

Gentiana hypericoides Gilg, Bot. Jahrb., 11: 312 (1896). Type: 

Ecuador, Lehman 652 1( K!). 
Gentianella hypericoides (Gilg) Fabris, Bol. Soc. Argent. Bot., 8: 184 

(1960). Calyx-lobes oblong-lanceolate; corolla c. 20-25 mm. 

Gentianella crossolaema (Wedd.) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana crossolaema Wedd., Chlor. And., 2: 55 (1859). Type: Peru, 
Weddel (P! - holotype). 

Gentianella cupiformis T. N. Ho & S. W. Liu, nom. nov. 

Gentiana campanuloides Gilg, Bot. Jahrb., 11: 320 (1916), non Willd. 

ex Roem. & Schult. (1820) (Roem. & Schult., Sy^r., 6: 184). Type: 

Argentina, Lorentz & Hieronymus (n.v.). 

Gentianella dasythamna (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana dasythamna Gilg, Bot. Jahrb., 54 (Beibl. 118): 63 (1917). 

Type: Bolivia, Larecaja, Mandon 363 ( P! - holotype, BM! K! - 

isotypes). 

Gentianella dissitifolia (Griseb.) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana dissitifolia Griseb., Gen. Sp. Gent., 229 (1839). Type: Peru, 
Dombey 394 (P! - lectotype designated here). 

Gentianella dombeyana (Wedd.) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana dombeyana Wedd., Chlor. And., 2: 62 (1845). Type: Peru, 
Dombey (P! - holotype). 

Gentianella ericothamna (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana ericothamna Gilg, Fedde, Rep. Nov. Sp., 2: 50 (1906). 
Type: Peru, Weberbauer 3381 (n.v.). 

Gentianella eurysepala (Gilg) Fabris ex T. N. Ho & S. W. Liu, comb, 
nov. 

Gentiana eurysepala Gilg, Bot. Jahrb., 50 (Beibl. Ill): 50 (1913). 
Type: Peru, Weberbauer 5616 (n.v.). 

Gentianella fiebrigii (Gilg) Holub, Folia Geobot. & Phytotax., Praha, 

2: 117(1967). 
Gentiana fiebrigii Gilg, Fedde, Rep. Nov. Sp. 2: 45 (1906). Type: 

Bolivia, K. Fiebrig 3187 (BM! K! - isotypes). 
Gentiana gynophora Gilg, Bot. Jahrb., IT. 305 (1896). Type: Bolivia, 

M. Bang 1231 (BM! K! - isotypes), syn. nov. 
There is no clear dividing hne between G. gynophora with long- 
petiolate basal leaves and scapose stem and G. fiebrigii with less 
long-petiolate basal leaves and scapose or foliate stem. G. gynophora 
is therefore not maintained as a distinct species. 

Gentianella fliipes (Cheeseman) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana filipes Cheeseman, Trans. Proc. N. Zeal. Inst., 28: 536 

(1896). Type: New Zealand, Mount Arthur, T. F. Cheeseman 

(BM! K! - isotypes). 

Gentianella florida (Griseb.) Holub, Folia Geobot. Phytotax., Praha, 
2: 117 ( 1967). Type: Argentina, Tucuman, Lorentz 310 (K!). 

Gentiana totorensis Gilg, Bot. Jahrb., 54(Beibl. 118): 84 (1917). 
Type: Bolivia, Herzog 2032 (n.v.), syn. nov. 

Gentianella fruticulosa (Domb. ex Wedd.) T. N. Ho & S. W. Liu, 
comb. nov. 

Gentiana fruticulosa Domb. ex Wedd., Chlor. And., 2: 71 (1839). 
Type: Peru, Dombey (P! - holotype). 

Gentianella gibbsii (Petrie) T. N. Ho & S. W. Liu, comb. nov. 

Gentiana gibbsii Petrie, Trans. Proc. N. Zeal. Inst., 49: 52. 1916 
(1917). Type: New Zealand, Stavart Island, F. G. Gibbs in herb. 
D. Petrie, W. 4709 (WELT! - holotype). 

Gentianella gracilifolia (Cheeseman) T. N. Ho & S. W. Liu, comb, 
nov. 

Gentiana gracilifolia Cheeseman, Man. N. Zeal. Fl. 1144 (1906). 
Type: New Zealand, Mount Arthur Plateau, F. G. Gibbs in herb. 
T. F. Cheeseman 137 (BM! K! - isotypes). 



NEW COMBINATIONS, NAMES AND NOTES ON GENTIANELLA 



63 



Gentianella graebneriana (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana graebneriana Gilg, Bot. Jahrb., 54(Beibl. 118): 24 (1917). 
Type: Peru, Weberbauer 6051 (n.v.). 

Gentianella grisebachii (Hook, f.) T. N. Ho, comb. nov. 

Gentiana grisebachii Hook, f., Ic. PL, t. 636 (1844). Type: New 
Zealand, Rotuari-Tongariro, J. C. Bidwili (K! - holotype). 

Gentianella ignea (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana ignea Gilg, Fedde, Rep. Nov. Sp., 2: 49 (1906). Type: Peru, 
Weberbauer 746 (n.v.). 

Gentianella krauseana (Gilg) Fabris ex T. N. Ho & S. W. Liu, comb, 
nov. 

Gentiana krauseana Gilg, Fedde, Rep. Nov. Sp., 2: 45 (1906). Type: 
Bolivia, K. Fiebrig 3187c (n.v.). 

Gentianella kuntzei (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana kuntzei Gilg, Bot. Jahrb., 22: 326 (May 1896). Type: 

Bolivia, O. Kuntze (n.v.). 
Gentiana cochabambensis Rusby, Mem. Torr. Bot. Club, 6: 76 (End 

1896). Type: Bolivia, M. Bang 1232 (BM! K! - isotypes). 

Gentianella kusnezowii (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana kusnezowii Gilg, Bot. Jahrb., 11: 325 (1896). Type: Bolivia, 
M. Bang 1230 (BM! K! - isotypes). 

Gentianella larecajensis (Gilg) T. N. Ho & S. W. Liu, comb. nov. 

Gentiana larecajensis Gilg, Bot. Jahrb., 54(Beibl. 118): 31 (1917). 
Types: Bolivia, Larecaja, Mandon 362 (P! - lectotype designated 
here, BM! K! - isolectotypes), 363 P. P. (BM! K!). 

Gentianella lilacina (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana lilacina Gilg, Fedde, Rep. Nov. Sp., 2: 40 (1906). Type: 
Peru, Weberbauer 3223 (n.v.). 

Gentianella lilacino-flavescens (Gilg) T. N. Ho & S. W. Liu, comb, 
nov. 

Gentiana lilacino-flavescens Gilg, Bot. Jahrb., 54(Beibl. 118): 38 
(1917). Type: Bolivia, Herzog 2114 (n.v.). 

Gentianella limoselloides ( Kunth ) Fabris, Bol. Soc. Argent. Bot., 8: 

166(1960). 
Gentianaana limoselloides Kunth, Nov. Gen., 3: 130 (1818). Types: 

Ecuador, Humboldt & Bonpland (P! - lectotype designated here), 

Bonpland 2266 {V\). 
Gentiana paludicola Gilg, Fedde, Rep. Nov. Sp., 2: 42 (1906). Type: 

Peru, Weberbauer 269A (n.v.), syn. nov. 
G. limoselloides is a widespread and variable species. It shows 
considerable variation especially in petiole length of the basal leaves 
and in cauline leaf shapes which are spathalate, oblong to lanceolate. 
Clearly, G. paludicola with lanceolate cauline leaves and long- 
petiolate basal leaves can not be maintained at any rank. 

Gentianella lineata (Kirk) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana lineata Kirk, Trans. Proc. N. Zeal. Inst. , 27: 334, t. 27. 1894 

(1895). Type: New Zealand, Hollows on the crest of the Longwool 

Range, T. Kirk (K! - isotype). 

Gentianella liniflora (Kunth) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana liniflora Kunth, Nov. Gen. Sp., 3: 171 (1818). Type: Peru, 
Humboldt & Bonpland ( P! - holotype). 

Gentianella lithophila (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana lithophila Gilg, Bot. Jahrb., 54(Beibl. 118): 39 (1917). 
Type: Bolivia, Herzog 2115 (n.v.) 

Gentianella lobbii (Gilg) T. N. Ho & S. W. Li u, comb. nov. 

Gentiana lobbii Gilg, Bot. Jahrb., 54 (Beibl. 118): 60 (1917). Type: 
Peru, Lobb (n.v.). 

Gentianella lurido-violacea (Gilg) Fabris ex T. N. Ho & S. W. Liu, 
comb. nov. 

Gentiana lurido-violacea Gilg, Fedde, Rep. Nov. Sp., 2: 37 (1906). 
Type: Peru, Weberbauer 7)159 (n.v.). 

Gentianella lythroides (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana lythroides Gilg, Bot. Jahrb., 54(Beibl. 118): 24 (1917). 
Type: Bolivia, Herzog 2229 (n.v.). 



Gentianella macrorriza (Gilg) Fabris ex T. N. Ho & S. W. Liu, comb, 
nov. 

Gentiana macrorriza Gilg, Bot. Jahrb., 54(Beibl. 118): 40 (1917). 
Type: Bolivia, Herzog 221 \ (n.v.). 

Gentianella magellanica (Gaudich.) Fabris in D. M. Moore, Vase., 

Fl. Falkland. Isl. 103 (1968). 
Gentiana magellanica Gaudich. in Freyc. Voy. Bot. 449 (1826). Type: 

lies Malouines, Gaudichaud (K! P! - isotypes). 
Gentiana pearcei Philippi, Anal. Univ. Santiago, 18: 65 (1861). Type: 

Chile, Philippi (K! - isotype), syn. nov. 
Gentiana valdiviana Philippi, Anal. Univ. Chil., 40: 206 (1895). 

Type: Chile, Philippi (K! - isotype), syn. nov. 
G. magellanica is a variable species in leaf, calyx-lobe shapes and 
corolla size. Leaves are oblong, ovate or lanceolate. Calyx-lobes are 
usually oblong, rarely ovate or lanceolate and all longer than the 
tube. The range of corolla size is (10-)13-18(-20) mm. The differ- 
ences in calyx-lobe shape and corolla size used to separate G. 
valdiviana and G. pearcei from G. magellanica are not of taxonomic 
significance. 

Gentianella mathewsii (Petrie) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana mathewsii Fetrie, Trans. Proc. N. Zeal. Inst., 44: 183. 1911 

(1912). Type: New Zealand, near Lake Harris, B. Petrie W. 4710 

(WELT! -holotype). 

Gentianella mesembrianthemoides (Gilg) T. N. Ho & S. W. Liu, 
comb. nov. 

Gentiana mesembrianthemoides Gilg, Fedde, Rep. Nov. Sp., 2: 41 
(1906). Type: Peru, Weberbauer 3303 (n.v.). 

Gentianella muscoides (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana muscoides Gilg, Fedde, Rep. Nov. Sp., 1: 35 (1906). Type: 
Peru, Weberbauer no. Ph. 96 (n.v.). 

Gentianella narcissoides (Gilg) T. N. Ho & S. W. Liu, comb, nov, 
Gentiana narcissoides Gilg, Bot. Jahrb., 54(Beibl. 118): 65 (1917). 
Type: Bolivia, Herzog 2414 (n.v.). 

Gentianella neomandonii (R. C. Foster) T. N. Ho & S. W. Liu, comb, 
nov. 

Gentiana neomandonii R. C. Foster, Rhodora, 56: 103 (1954), based 
on Gentiana mandonii Gilg, Bot. Jahrb., 54(Beibl. 118): 37 (1917), 
non Rusby (1896) {Mem. Torr. Bot. Club, 6: 80). Type: Bolivia, 
Mandon 363a (P! - holotype, BM! K! - isotypes). 

Gentianella odontosepala (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana odontosepala Gilg, Fedde, Rep. Nov. Sp., 2: 48 (1906). 
Type: Bolivia, M. Bang 261 \ (n.v.). 

Gentianella oreosilena (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana oreosilena Gilg, Fedde, Rep. Nov. Sp., 2: 40 (1906). Type: 
Peru, Weberbauer 4288 (n.v.). 

Gentianella orobanchoides (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana orobanchoides Gilg, Bol. Jahrb., 11: 333 (1896). Type: 

Bolivia, Mandon 366 (P! - lectotype designated here, BM! K! - 

isolectotypes). 

Gentianella pachystemon (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana pachystemon Gilg, Bot. Jahrb., 54(Beibl. 118): 46 (1917). 
Syntypes: Peru, Stuebel 51, 52 (all n.v.). 

Gentianella palcana (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana palcana Gilg, Bot. Jahrb., 54(Beibl. 118): 47 (1917). 
Syntypes: Bolivia, Herzog 2116 (n.v.), Stuebel 46c (n.v.). 

Gentianella pallido-lilacina (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana pallido-lilacina Gilg, Bot. Jahrb., 54(Beibl. 118): 58 (1917). 
Type: Bolivia, Herzog 2028 (n.v.). 

Gentianella parviflora (Griseb. ) T. N. Ho, stat. et comb. nov. 

Gentiana coerulescens Gill, ex Wedd. var. parviflora Griseb., Symb. 
Fl. Argent., 237 (1879). Type: Argentina, Hieronymus 464 (K!). 

Gentianella peruviana (Griseb.) Fabris, Bol. Soc. Argent. Bot., 7: 93 

(1958). 
Gentiana limoselloides Kunth var. peruviana Griseb., Gen. Sp. 

Gent., 215 (1838). Type: Peru, Weddell 4445 (P! - Type). 
Gentiana peruviana (Griseb.) Gilg, Bot. Jahrb. 11: 304 (1896), non 



64 

Lamarck (1786) {Lamarck, Encycl. Meth. 2: 642). 
G. peruviana is a perennial dwarf herb characterized by a rather 
fleshy tap-root, developed, oblong to oblanceolate basal leaves and 
oblong, ovate-oblong to obovate calyx-lobes which are as long as, 
slightly shorter or longer than the tube. It includes four species (G. 
peruviana, G. hieronymii, G. boliviana and G. lobelioides) which are 
better treated as the following two varieties. The difference between 
var. peruviana and var. boliviana is only of corolla size: larger flowers 
(16-17 mm) of the former and smaller flowers (11-13 mm) of the 
latter. 

var. peruviana 

Gentiana hieronymii Gilg, Bot. Jahrb., 22: 305 (1896). Type: Argen- 
tina, Lorentz & Hieronymus 15 (K! - isotype), syn. nov. 

Gentianella hieronymii (Gilg) Fabris, Rev. Invest. Agric, Buenos 
Aires, 11: 396 (1958), in adnot. 

var. boliviana (Pax) T. N. Ho, stat. et comb. nov. 

Gentiana boliviana Pax, Fedde, Rep. Nov. Sp., 7: 243 (1909). Type: 

Bolivia. Buchtien 1482 (n.v.). 
Gentiana lobelioides Gilg, Bot. Jahrb., 54(Beibl. 118): 30 (1917). 

Type:Peru, Weberbauer 955 (n.v.), syn. nov. 

Gentianella petrophila (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana petrophila Gilg, Fedde, Rep. Nov. Sp., 2: 42 (1906). Type: 
Peru, Weberbauer 2562 (n.v.). 

Gentianella pilgeriana (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana pilgeriana Gilg, Bot. Jahrb., 54(Beibl.ll8): 42 (1917). 
Type: Bolivia, Herzog 2410 (n.v.) 

Gentianella poculifera (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana poculifera Gilg, Bot. Jahrb., 50(Beibl. 118): 48 (1913). 
Type: Peru, Weberbauer (n.v.) 

Gentianella porphyrantha (Gilg) Fabris ex T. N. Ho & S. W. Liu, 
comb. nov. 

Gentiana porphyrantha Gilg, Fedde, Rep. Nov. Sp., 2: 39 (1906). 
Type: Peru, Weberbauer 2803 (n.v.). 

Gentianella potamophila (Gilg) Fabris ex T. N. Ho , comb. nov. 

Gentiana potamophila Gilg, Bot. Jahrb., 54(Beibl. 118): 74 (1917). 
Type: Peru, Weberbauer 6907 (n.v.). 

Gentianella primuloides (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana vaginalis Griseb. ex Wedd. Chlor. And., 2: 53 (1859), non 

Griseb. (1838) (Gen. Sp. Gent., 2\5). 
Gentiana primuloides Gilg, Bot. Jahrb., 54(Beibl. 118): 37 (1917). 

Types: Peru, techier 2002 (P! - lectotype designated here); 

Bolivia, M. Bang 1888 (BM! K!), 1889 (BM! K!), O. Kuntze 

(n.v.). Bock in herb. Herzog 2480c (n.v.), Herzog 2081 (n.v.). 

Gentianella pseudolycopodium (Gilg) T. N. Ho & S. W. Liu, comb, 
nov. 

Gentiana pseudolycopodium Gilg, Fedde, Rep. Nov. Sp., 2: 38 
(1906). Type: Peru, Huamalies, Weberbauer 3353 (n.v.). 

Gentianella pulla (Griseb.) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana pulla Griseb., Goett. Abh., 19: 209 (1874). Type: Argen- 
tina, Lorentz 111) (K! - isotype). 

Gentianella punicea (Wedd.) Holub, Folia. Geobot. Phytotax., 

Praha, 2: 118(1967). 
Gentiana punicea Wedd., Chlor. And., 2: 70 (1859). Types: Bolivia, 

Mandon 364 (K! BM!); Peru, Weddell 4741 (P! - lectotype 

designated here). 
Gentiana dolichantha Gilg, Torrey, 5: 109 (1909). Syntype: Bolivia, 

Lobb (n.v.), R. S. Wiliams 2489 (K! BM!), syn. nov. 
Gentiana purpureiflora Gilg, Bot. Jahrb. 54 (Beibl. 118): 65 (1917). 

Type: Bolivia, Herzog 2168 (n.v.), syn. nov. 
The authors could find no distinction between G. dolichantha and G. 
punicea in any respect. G. purpureiflora is also identical to G. 
punicea in habit, leaf shape, corolla size as well as the character of the 
calyx-lobes. It seems that G. purpureiflora is supposed to differ from 
G. punicea in its corolla-lobes which are as long as the tube. 
However, there is a complete range of intermediates from short to 
long corolla-lobes in G. punicea. Therefore, G. dolichantha and G. 
purpureiflora are all reduced to synonyms. 



T.-N. HO AND S.-W. LIU 

Gentianella raimondiana (Wedd.) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana raimondiana Wedd., Chlor. And., 2: 310 (1861). Type: 
Peru, Raimondi (n.v.). 

Gentianella roseo-lilacina (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana roseo-lilacina Gilg, Fedde, Rep. Nov. Sp., 2: 35 (1906). 
Type: Peru, Weberbauer 2952 (n.v.). 

Gentianella sancti-mathacii (R. C. Foster) T. N. Ho & S. W. Liu, 
comb. nov. 

Gentiana sancti-mathacii R. C. Foster, Rhodora, 56: 103 (1954), 
based on Gentiana praticola Gilg, Bot. Jahrb., 54(Beibl. 118): 37 
(1917), non Franchet (1896). {Bull. Soc. Bot. Fr., 43: 489). Type: 
Bolivia, Herzog 1977 (n.v.). 

Gentianella sandiensis (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana sandiensis Gilg, Fedde, Rep. Nov. Sp., 2: 36 (1906). 

Syntypes: Bolivia, Weberbauer 1016 (n.v.); Peru, Weberbauer 352, 

445, 917a, 2593 (all n.v.), Poeppig (n.v.). 

Gentianella saxicola (Griseb.) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana saxicola Griseb., Gen. Sp. Gent., 216 (1838). Type: Peru, 

Andes, Mathews (K! - isotype). 
Gentiana vaginalia Griseb., Gen. Sp. Gent., 215 (1838). Type: Peru, 

Pasco, Cruikshanks (K!), syn. nov. 
The type of G. vaginalia is identical to the type of G. saxicola in all 
respects, e. g. dwarf herb, crowded leaves forming a rosette, leaves 
ovate or triangular, calyx-lobes all papillose on upper surfaces and 
distinctly ciliate on margins. G. vaginalia is therefore not maintained 
here. 

Gentianella scarlatiflora (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana scarlatiflora Gilg, Bot. Jahrb., 50 (Beibl. Ill): 49 (1913). 
Syntypes: Peru, Lobb (n.v.), Weberbauer 5836 (n.v.). 

Gentianella scarlatina (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana scarlatina Gilg, Fedde, Rep. Nov. Sp., 2: 36 (1906). Type: 
Peru, Sandia, Weberbauer 1047 (n.v.). 

Gentianella scarlatino-striata (Gilg) T. N. Ho & S. W. Liu, comb, 
nov. 

Gentiana scarlatino-striata Gilg, Bot. Jahrb., 54(Beibl. 118): 67 
(1917). Type: Peru, Weberbauer 6621 (n.v.). 

Gentianella scopulorum (Wedd.) Fabris ex T. N. Ho & S. W. Liu, 
comb. nov. 

Gentiana scopulorum Wedd., Chlor. And., 32: 67 (1859), nee Tide- 
str. (1925) (Contr. U.S. Nat. Herb., 26: 416). Type: Bolivia, 
Weddell 3928 (P! - holotype). 

Gentianella silenoides (Gilg) Fabris in A. L. Cabrera, Fl. Prov. Jujuy 

{Inst. Nac. Teen. Agropec, 13), 8: 68 (1983). 
Gentiana silenoides Gilg, Bot. Jahrb., 22: 319 (1896). Type: Bolivia, 

Lorentz & Hieronymous 878 P. P. (n.v.). 

var. silenoides 

Flowers in a lax cyme; corolla 16-18 mm. 

var. striaticalyx (Gilg) Ho, stat. et comb. nov. 

Gentiana striaticalyx Gilg, Bot. Jahrb., 54(Beibl. 118): 56 (1917). 

Type: Bolivia, Herzog20A6h. (n.v.). 
Gentiana anthosphaera Gilg, Fedde, Rep. Nov. Sp., 2: 46 (1906). 

Type: Bolivia, Fiebrig 2246 (K! P! - isotypes), syn. nov. 
Gentiana herzogii Gilg, Bot. Jahrb., 54 (Beibl. 118): 57 (1917). Type: 

Bolivia, Herzog 2018 (n.v.), syn. nov. 
Gentianella anthosphaera (Gilg) Holub, Folia Geobot. & Phytotax., 

Praha, 2: 116 (1967); Fabris in A. L. Cabrera, Fl. Prov. Jujuy {Inst. 

Nac. Teen. Agropec, 13), 8: 77 (1983), comb, superfl. 
Flowers in a lax cyme; corolla 20-30 mm. 

var. inaequicalyx (Gilg) Ho, stat. et comb. nov. 

Gentiana inaequicalyx Gilg, Bot. Jahrb., 22: 324 (1896). Syntypes: 

Bolivia, Mandon 365 (Ki), M. Bang 1143 (K!), O. Kuntze (n.v.). 
Flowers in a dense umbel or subcapitulum; corolla 20-30(-35) mm. 

Gentianella serotina (Cockayne) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana serotina Cockayne, Trans. Proc. N. Zeal. Inst., 47: 113. 
1914 (1915). Type: New Zealand, W. 4724 (n.v.). 

Gentianella setipes (Gilg) T. N. Ho & S. W. Liu, comb. nov. 



NEW COMBINATIONS, NAMES AND NOTES ON GENTIANELLA 



65 



Gentiana setipes Gilg, Bot. Jahrb., 54(Beibl. 118): 43 (1917). Type: 
Peru, Weberbauer 6322 (n.v.). 

Gentianella soratensis (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana soratensis Gilg, Bot. Jahrb., 22: 332 (1896). Type: Bolivia, 
Rusby 675 (P! - holotype, BM! K! - isotypes). 

Gentianella speciosissima (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana speciosissima Gilg, Bot. Jahrb., 22: 325 (1896). Type: Peru, 
Stuebel 2Ah, 25 {a\\ n.v.). 

Gentianella spedenii (Petrie) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana spedenii Petrie, Trans. Proc. N. Zeal. Inst., 56: 14 (1926). 

Type: New Zealand, Princess Range, James Speden (WELT! - 

holotype). 

Gentianella spenceri (Kirk) T. N. Ho & S. W. Liu, comb. nov. 

Gentiana spenceri T . Kirk, Trans. Proc. N. Zeal. Inst., 27: 335, t. 27 
A, B. 1894 (1895). Type: New Zealand, Mount Rochfort, F. H. 
Spencer, W. 4711 (n.v.). 

Gentianella stellarioides (Gilg) Fabris, Bol. Soc. Argent. Bot. , 8: 180 

(1960). 
Gentiana stellarioides Gilg, Bot. Jahrb., 54(Beibl. 118): 79 (1917). 

Types: Ecuador, Spruce (K! - lectotype designated here), Jameson 

(K!). 

var. stellarioides 

var. androtricha ( Gilg) T. N. Ho, stat. et comb. nov. 

Gentiana androtricha Gilg, Bot. Jahrb., 54(Beibl. 118): 79 (1917). 

Type: Ecuador, Spruce 6050 (K!). 
This variety is very similar to var. stellarioides in having linear to 
linear-lanceolate calyx-lobes, very long pedicels and large flowers 
(17-20 X 15-20 mm), but is distinguished by basal leaves with long 
petioles and elliptic cauline leaves. 

Gentianella stenosepala (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana stenosepala Gilg, Bot. Jahrb., 22: 331 (1896). Type: 
Bolivia, O. Kuntze (n.v.). 

Gentianella stricticaulis (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana stricticaulis Gilg, Bot. Jahrb., 54(Beibl. 118): 62 (1917). 
Type: Peru, Weberbauer 6097 (n.v.). 

Gentianella stuebelii (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana stuebelii Gilg, Bot. Jahrb., 22: 317 (1916). Type: Peru, 
Stuebel35{{n.v.). 

Gentianella tarapacana (Gilg) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana tarapacana Gilg, Bot. Jahrb., 22: 305 (1896). Type: Chile, 
Philippi (K! - isotype). 

Gentianella tenuifolia (Petrie) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana tenuifolia Petrie, Trans. Proc. N. Zeal. Inst., 45: 270. 1912 

(1913). Type: New Zealand, near Lyell, S. W. Nelson in herb. B. 

Petrie, W. Townson, W. 4721 (WELT! - holotype). 

Gentianella tereticaulis (Petrie) T. N. Ho & S. W. Liu, comb. nov. 
Gentiana tereticaulis Petrie, Trans. Proc. N. Zeal. Inst., 49: 51. 1916 

(1917). Type: New Zealand, Lake Harries, Routeburn Valley, W. 

Petrie (WELT! - holotype). 

Gentianella thiosphaera (Gilg) Holub, Folia Geobot. Phytotax., 

Praha, 2: 118(1967). 
Gentiana thiosphaera Gilg, Fedde, Rep. Nov. Sp.,2: 46 (1906). Type: 

S. Bolivia, K. Fiebrig 3156 (P! - holotype, BM! K! - isotypes). 

var. thiosphaera 

Gentiana comarapana Gilg, Bot. Jahrb., 54(Beibl. 118): 82 (1917). 

Type: Bolivia, Herzog 1914 (n.v.), syn. nov. 
G. comarapana was originally described as having lanceolate calyx- 
lobes which are as long as the tube, and thus differs from G. 
thiosphaera. The calyx-lobes of both species are lanceolate, linear- 
lanceolate to linear and as long as to much longer than the tube. G. 



comarapana is therefore not maintained here. 

var. macroclada (Gilg) Ho, stat. et comb. nov. 

Gentiana macroclada Gilg, Fedde, Rep. Nov. Sp., 2: 47 (1906). Type: 

Bolivia, K. Fiebrig 2654a. (n. v.). 
This variety is very similar to var. thiosphaera and is distinguished 
only by smaller flowers (usually 12-14 mm). 

Gentianella townsonii (Cheeseman) T. N. Ho & S. W. Liu, comb, 
nov. 

Gentiana townsonii Cheeseman, Man. N. Zeal. FL, 450 (1906). 
Syntypes: New Zealand, Mount Rochfort, W. Townson 288 (BM! 
K!), 389(BM!K!). 

Gentianella tristicha (Gilg) Fabris ex T. N. Ho & S. W. Liu, comb, 
nov. 

Gentiana tristicha Gilg, Fedde, Rep. Nov. Sp., 2: 39 (1906). Type: 
Peru, Weberbauer 2933 (n.v.). 

Gentianella vernicosa (Cheeseman) T. N. Ho & S. W. Liu, comb, 
nov. 

Gentiana vernicosa Cheeseman, Man. N. Zeal. FL, 1145 (1906). 
Type: New Zealand, Mount Lockett, F. G. Gibbs (n.v.). 



Acknowledgements. We are grateful to the directors and staff of 
the herbaria of The Natural History Museum, Royal Botanic Gar- 
dens, Kew, Museum National d'Histore Naturelle, Paris and the 
National Museum of New Zealand, for permission to consult their 
collections and for the loan of specimens. We are much indebted to 
Mr Roy Vickery, Miss Sally Bidgood and Mr Thierry Deroin for their 
help in providing most of the herbarium material. Thanks are also 
due to Dr Mike Gilbert for useful suggestions in the preparation of 
this paper. 



REFERENCES 



Allan, H. H., 196L Gentiana. In Flora of New Zealand 1: 764-779. Wellington. 
Fabris, H. A., 1953. Sinopsis preliminar de las Gentianaceae Argentinas, Bol. 

Soc. Argent. Bot., 4: 232-259. 
, 1955. Nuevas especies de 'Gentianella' del Peru, Bol. Soc. Argent. Bot., 

6(1): 45-50. 
, 1958. Notas Sobre Gentianella del Peru. Bol. Soc. Argent. Bot., 7(2): 

86-93. 
, 1959. Sobre la identid de dos species Sudamericanae de Gentianella, Bol. 

Soc. Argent. Bot., 8(1): 24-25. 
, 1960. El Genero Gentianella en Ecuador, Bol. Soc. Argent. So?., 8(3-4): 

160-191. 
Gilg, E., 1896. Beitrage zur Kenntnis der Gentianaceae 1. Bot. Jahrb.. 22: 

301-347. 
, 1906. Beitrage zur Kenntnis der Gentianaceae III. Gentianaceae, 

Andinae, Fedde, Rep. Nov. Sp., 2: 34-52. 
, 1917. Monographische Zusammenstellung der Gentiana-Arten Sud- 

Amerikas, Bot. Jahrb., 54(Beib. 118): 4-89. 
Gillet, J. N., 1957. A. revision of the north American species of Gentianella 

Moench, >l«/i. Missouri Bot. Gard., 44(3): 195-262. 
Grisebach, A. H. R., 1838 (1839). Genera et species Gentianearum. Stuttgart. 
Holub, J., 1967. Neue namen innerhalb der Gattungen Gentianella Moench, 

Gentianopsis Ma and Comastoma (Wettst.) Toyokuni, Folia Geobot, Phyto- 
tax. Praha, 2: 115-120. 
, 1968. Einige neue nomenklatorische Kombinationen innerhalb der 

Gentianinae, Folia Geobot. Phytotax. Praha, 3: 217-218. 

, 1983. A brief note on Slovak taxa of Gentianella, Preslia, 55(4): 371-373. 

Pringle, J. S., 1981, Nomenclatural transfers and taxonomic notes on some 

South American Gentianaceae, Phytologia, 48(4): 281-285. 
, 1987. A new species and taxonomic notes on Gentianella (Gentianaceae) 

in South America, Sida, 11(4): 357-369. 



Bull. nat. Hist. Mus. Lond. (Bot.) 23(2): 67-70 



Issued 25 November 1993 



Studies in Hypericum: validation of new 
names 

NORMAN K.B. ROBSON 

do Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD 

CONTENTS 



Introduction 67 

Section 20. Myriandra (Spach) R. Keller 67 

Section 23. Triadenioides Jaub.& Spach 68 

Section 25. Adenotrias (Jaub.& Spach) R. Keller 68 

Section 27. Adenosepalum Spach 69 

References 70 



Synopsis. The new names in Part 6 of 'Studies in the genus Hypericum L. (Guttiferae)' are validated in advance of 
publication of the main work. They are:- new taxa (tax. nov.): H. hypericoides (L.) Crantz subsp. prostratum N. 
Robson (Sect. 2Q. Myriandra), H. fieriense N. Robson (Sect. 23. Triadenioides), subsects. Aethiopica N. Robson, 
Pubescentes N. Robson and H. collenettiae N. Robson (Sect. 27. Adenosepalum); new combinations (comb, et stat. 
nov.): H.nitidum subsp. cubense (Turcz.) N. Robson, H. nitidum subsp. exile (P. Adams) N. Robson,//. aegypticum 
L. subsp. maroccanum (Pau) N. Robson, //. annulatum Moris subsp. intermedium (Steudel ex A. Rich.) N. Robson, 
//. annulatum subsp. afromontanum (Bullock) N. Robson. 



INTRODUCTION 



Part 6 of 'Studies in the genus Hypericum L. (Guttiferae)', a 
monographic series that is intended to cover the whole genus 
(Robson 1977, 1981, 1985, 1987, 1990), will contain accounts 
and analyses of Sections 20-28. As this part will not be 
published before 1994, it is necessary to validate in advance 
of publication the new names that will appear in it. 

Section 20. MYRIANDRA (Spach) R. Keller 

Hypericum nitidum Lam., Encycl. 4:160 (1797). 

When the populations from Cuba and Belize in the H. 
nitidum group are considered along with those in the U.S.A., 
they can be divided into three subspecies: a) Cuba and Belize 
(subsp. cubense), which is related to the Cuban H. limosum 
Griseb.; b) south-eastern U.S.A. (subsp. nitidum); and c) 
western Cuba and north-western Florida (subsp. exile), which 
is related to H. brachyphyllum (Spach) Steudel from south- 
eastern U.S.A. 



H. nitidum subsp. nitidum 

Leaves subcoriaceous, apex obtuse to rounded-apiculate, 
margin loosely inrolled leaving lower lamina partly exposed. 
Sepals obtuse to shortly apiculate. Capsule cylindric. 

U.S.A. (south-eastern Alabama to southern N. Carolina). 

©The Natural History Museum, 1993 



H. nitidum subsp. cubense (Turcz.) N. Robson, comb, 
et stat. nov. 

H. cubense Turcz. in Bull. Soc. Nat. Moscou 31(1): 384 

(1858). 

H. fasciculatum sensu Alain in Leon & Alain, Fl. Cuba 3:317 

(1953) pro parte. 

Leaves coriaceous, apex rounded-apiculate to rounded, mar- 
gin tightly inrolled leaving only midrib exposed. Sepals 
obtuse to rounded-apiculate. Capsule cylindric to rarely 
ovoid-conic. 

Cuba: (Oriente, Las Villas, Isla de Pinos), Belize (El Cayo). 

Hypericum nitidum Lam. subsp. exile (P. Adams) N. 
Robson, comb, et stat. nov. 

H. galioides var. cubense Griseb., Cat. PI. Cuba: 39 (1866), 

non H. cubense Turcz. (1858). 
H. galioides var. axillare sensu Griseb., loc. cit., pro parte 

(1866). 
H. galioides sensu Sauvalle, Fl. Cubana: 8 (1868). 
H. fasciculatum sensu Alain in Leon & Alain, Fl. Cuba 3: 317 

(1953) pro parte excl. typum. 
H. exile P. Adams in Contr. Gray Herb. Harv. no. 189: 33 

(1962). 

Leaves chartaceous, apex acute to long-acuminate, margin 
tightly inrolled leaving only midrib exposed. Sepals acute to 
long-acuminate. Capsule cylindric to narrowly conic. 

U.S.A. (north-western Florida), Cuba (Pinar del Rio, Isla de 
Pinos). 



68 



N.K.B. ROBSON 



Hypericum hypericoides (L.) Crantz, Inst, rei herb. 
2:520 (1776). 

Ascyrum hypericoides L., Sp. pi. :789 (1753) excl. syn. Hort. 
Cliff, et Plukenet., 2nd ed.: 1108 (1763) excl. syn. 
Plukenet. Type: Hispaniola, Hypericoides frutescens erecta, 
flore luteo Plumier, Nov. pi. amer.: t.7 (1703), lectotype 
(Robson, 1980:272). 

Plumier (1703) distinguished two species from Hispaniola in 
his new genus Hypericoides: H. frutescens, erecta, flore luteo 
and H. frutescens, humi-fusa, flore luteo. The erect species is 
not distinguishable taxonomically from Hypericum hyperi- 
coides subsp. hypericoides as it is known in the other islands 
of the Greater Antilles, the Bahamas, Bermuda and the 
mainland (eastern N. America, eastern Mexico to Honduras 
Republic); but the other represents a taxon that has appar- 
ently evolved within Hispaniola, occurring at high altitudes in 
the Dominican Republic. The occurrence of a few somewhat 
intermediate specimens in the region between 1600 and 
2000 m, where it co-exists with the typical form, indicates that 
the appropriate rank for this taxon is subspecies. 



Hypericum hypericoides subsp. prostratum N. 
Robson, subsp. nov. 

Hypericoides frutescens, humi-fusa, flore luteo Plumier, Nov. 

pi. amer.: 52(1703). 
Ascyrum foliis lanceolato-linearibus, biglandulosis, ramidiffu- 

sis Burman, PI. amer.: 146, t.l52 f.2 (1758). 

A subsp. hypericoides habitu prostrato, folium minoribus 
anguste oblongis vel oblongo-spathulatis, differt. 

Type: Dominican Republic, San Juan, Sabana Nueva, Cor- 
dillera Central N. of Rio Arriba del Norte, 1950 m, 
17-20. ix. 1944, R.A. & E.S. Howard 9080 (BM!, holotype; 
GH!, MICH!, NY!, US!, isotypes). 

Plant prostrate, with stems ? numerous, radiating and 

branching, forming mats. Leaves 3-8(-10) X 1-2.5 mm, 

narrowly oblong to oblong-spathulate. Inflorescence- 
branching pseudo-dichotomous. 

Open Pinus forest, grassland and open slopes, 
(160(>-)1800-2900 m. 

Dominican Republic (La Vega, Santiago, San Juan, Peravia). 



Section 23. TRIADENIOIDES Jaub. & Spach 

Socotra, an island of endemics, is critical for the understand- 
ing of the evolution of Hypericum. It is already known to 
contain four endemic species, H. balfourii N. Robson and H. 
socotranum Good (Sect. 1. Campylosporus), the latter with 
two subspecies (Robson 1985), H. scopulorum Balf. f. and H. 
tortuosum Balf. f. (Sect. 23. Triadenioides). To these must be 
added a new species based on one collection from the 
Hagghiher Mountains by Smith and Lavranos. It is clearly 
near H. scopulorum but is more woody and larger in all its 
parts and has petiolate leaves, and the lower leaf surface, 
petiole and young stems are covered with a puberulous 
indumentum. 



Hypericum fieriense N. Robson, sp. nov. 

H. scopulorum Balf. f. affinis, sed caulibus crassioribus 
lignosioribus, foliorum lamina ovata subtus cum petiolo et 
ramis junioribus puberula, inflorescentia 3-5-florata, floribus 
maioribus sepalis crassioribus, capsulis coriaceis valvis leviter 
angustissime vittatis fere laevibus, inter alia differt. 

Type: Socotra, Hagghiher Mountains(12°35'N, 54°03'E), 
below Fieri peaks, 1350 m, 21. iv. 1967, Smith & Lavranos 475 
(K!, holotype & isotype). 

'Low scrub among Dracaena cinnabari trees', 1350 m. 

Socotra (Hagghiher Mts). 

Section 25. ADENOTRIAS (Jaub. & Spach) R. 
Keller 



Hypericum aegypticum L., Sp. PL: 784 (1753). 

H.aegypticum comprises a series of disjunct populations 
forming a morphological reduction trend from south 
Morocco to Crete and Cyrenaica (not Egypt). This trend is 
almost continuous, but it is possible to recognize three, rather 
poorly differentiated subspecies. Only essential synonymy is 
given here. 

Hypericum aegypticum L. subsp. maroccanum (Pau) 
N. Robson, stat. nov. 

H. aegypicum var. maroccanum Pau in Cavanillesia 4: 157 
(1932) ['maroccana'] in Spanish; Maire in Bull. Soc. Hist, 
nat. Afr. N. 24: 206 (1933), in Latin. 

Plant erect, (0.15-)0.3-2 m tall, with branches erect to 
ascending. Leaves sessile, plane; lamina (7-)9-18 X 
(2-)3-4 mm, narrowly elliptic or narrowly oblong-elliptic, 
acute. Sepals 5-6 mm long. Petals 10-12(14?) mm long. 

Morocco (south-west), Algeria (southern Atlas Mts). 

H. aegypticum L. subsp. webbii (Spach) N. Robson, 
comb. et. stat. nov. 

Triadenia webbii Spach in Annls Sci. nat. (Bot.) II, 5: 174, t. 
5A (1836), Hist. nat. veg. Phan. 5: 372 (1836). 

Plant erect to loosely spreading, 0.04—0.4 m tall, with 
branches erect or usually spreading and often tortuous, 
forming bushes up to 1 m across. Leaves subsessile to shortly 
(c. 0.3 mm) petiolate, plane or subcucuUate; lamina 4-10 X 
1.5-3 mm, narrowly oblong to broadly elliptic; acute to 
obtuse. Sepals 5-6 mm long. Petals 8-14 mm long. 

Lampedusa, Malta, Sardinia, Greece (Ionian Islands, west- 
ern Peloponnisos), Crete. 

H. aegypticum L. subsp. aegypticum 

Plant spreading 0.05-0.18 m tall, with branches ± spreading 
and tortuous, forming low bushes. Leaves shortly ± 
(0.4-0.5 mm) petiolate, always (?) ± incurved-cucullate; 
lamina 3-6 X 1-2 mm, narrowly oblong to broadly elliptic, 
acute. Sepals 3.5-5 mm long. Petals 6.5-9 mm long. 

Libya (Cyrenaica-Jebel el Akhdar). 



STUDIES IN HYPERICUM 



69 



Section 27. ADENOSEPALUM Spach 

When Sect. Adenosepalum (sensu Robson, 1977) has been 
'purified' by the removal of the tropical Asian species(//. 
elodeoides group) to Sect. 9. Hypericum sensu lato and three 
Turkish species {H. huber-morathii N. Robson, H. minutum 
P.H. Davis & Poulter, H. formosissimum Takht.) to Sect. 12. 
Origanifolia, the remaining species form a natural group 
distributed over most of Africa, Macaronesia, Europe, Medi- 
terranean Asia and western Arabia. It can be divided into 
four subsections, as follows: 

1. Subsect. Aethiopica N. Robson, subsect. nov. Planta 
omnino glabra. Folia libera. Bracteae bracteolaeque haud 
glanduloso-auriculatae. Typus: H. aethiopicum Thunb. 

2. Subsect. Pubescentes N. Robson, subsect. nov. Planta 
usque ad sepala vel ad partem inferiorem inflorescentiae 
vel rare ad basin inflorescentiae indumentum ferens. Folia 
libera. Bracteae bracteolaeque haud glanduloso- 
auriculatae. Typus: H. pubescens Boiss. 

3. Subsect. Caprifolia N. Robson, subsect. nov. Planta usque 
ad basin inflorescentiae indumentum ferens. Folia inter- 
dum inferiora excepta binatim conjuncta. Bracteae 
bracteolaeque interdum glanduloso-auriculatae. Typus: 
H. caprifolium Boiss. 

4. Subsect. Adenosepalum. Planta usque ad basin inflores- 
centiae indumentum ferens vel rare caulibus vel foliis vel 
omnino glabra. Folia libera. Bracteae bracteolaeque per- 
saepe glanduloso-auriculatae. Typus: H. montanum L. 



treated as conspecific with the central Balkan H. degenii 
Bornm. and the Ethiopian and East African H. intermedium 
Steudel ex A. Rich. (Milne-Redhead, 1953 a,b; Robson, 
1958, 1968; etc.). 

Further study of this variable species and its unusual discon- 
tinuous distribution has shown a) that the widely separate 
populations can be differentiated as three subspecies and b) 
that the Mt. Elgon endemic H. afromontanum Bullock is no 
more than a high-altitude form of the East African subspe- 
cies. The Ethiopian/Arabian subsp. intermedium is morpho- 
logically nearest to H. montanum L., the sister species of H. 
annulatum. 

H. annulatum Moris subsp. annulatum 

H. perfoliatum var. annulatum (Moris) Fiori in Fiori & 
Paoletti, Fl. Anal. It. 1: 389 (1898), Nuovo Fl. Anal. It. 1: 
524 (1924). 

H. degenii Bornm. in Magyar Bot. Lap. 9:90 (1910). 

Stem without red or black glands, densely shortly pubescent. 
Leaves without laminar black glands, densely shortly pubes- 
cent. Sepals long- to short-glandular-ciliate (cilia shorter 
than to two or more times as long as glands), with laminar 
glands all pale. Petals without punctiform laminar black 
glands, not tinged red in bud. 

Sardinia, [Yugoslavia] (southern Serbia, Macedonia), north- 
ern Albania, Bulgaria, northern Greece. 



Hypericum coUenettiae N. Robson, sp. nov. 

H. sp. aff. sinaicum sensu Collenette, ///. Guide Fls Saudi 
Arabia: 262 + photos (1985). 

H. somalensi N. Robson affinis, sed indumento breviori, 
caulibus internodiis plerumque foliis brevioribus, foliis angus- 
tioribus, inflorescentia paucioriflora laxiore, floribus maiori- 
bus, sepalis petalisque glandulis nigris laminaribus ornatis, 
inter alia differt. 

Type: Saudi Arabia, Asir, Taif-Abha road 82 km S. of 
Baljurshi, Wadi Mahra, c.1800 m, 5.viii.l982 (fr), Collenette 
3752 (BM!, holotype; K!, isotype). Collenette 1401 (K) is 
another collection (16. iv. 1979) from the same population. 

Shady rock crevices, c.1800 m. 

Saudi Arabia (Asir). 

This apparently solitary population is intermediate in mor- 
phology and distribution between H. somaliense N. Robson 
(N. Somalia) and H. sinaicum Hochst. ex Boiss. (Sinai and 
adjacent Saudi Arabia) but distinct from both these species. 
According to Mrs. Collenette, its habitat is under some 
threat. An earlier specimen from the same area (between 
Baljurshi and Abha) has recently come to light and may 
represent a second population:- Bashwat, 9.viii. 1975 (fl. & 
fr.), A. El-Sheikh in Herb. KSUH 1067 (KSUH). 

Hypericum annulatum Moris, Stirp. sard, elench.: 9 
(1827). 

Although originally regarded as endemic to Monte Santa 
Vittoria esterzili in Sardinia, this species has subsequently 
been found in another locality in that island (Nodu 'e 
Littipori) (Arrigoni et al. 1973). Meanwhile it had been 



H. annulatum Moris subsp. intermedium (Steudel ex 
A. Rich.) N. Robson, comb, et stat. nov. 

H. intermedium Steudel ex A. Rich., Tent. Fl. Abyss. 1: 95 
(1847). 

H. intermedium forma obtusifolium R. Keller ex Moggi & 
Pisacchi in Webbia 22: 272 (1967) in synon. 

H. annulatum sensu Cufod. in Bull. Jard. bot. Etat Brux. 29, 
Suppl.: 588 (1959); Moggi & Pisacchi in Webbia 11: 111 
(1967) pro parte; Collenette, ///. Guide Fls Saudi Arabia: 
261 + photo (1985), pro parte excl. typum et spec. cit. ex 
Harar. 

Stem without or rarely with few black glands, ± sparsely 
puberulous to glabrous. Leaves without laminar black glands, 
± sparsely and very shortly pubescent to puberulous or 
glabrous. Sepals short- to long-glandular-ciliate (cilia shorter 
than to two or more times as long as glands), occasionally 
with some laminar glands black. Petals with few (rarely more 
numerous) punctiform laminar black glands, rarely red- 
veined in bud. 

Saudi Arabia (Asir), Sudan Republic (southern Red Sea 
Hills), northern Ethiopia (Eritraea to L. Tana and northern 
Shoa). 

H. annulatum Moris subsp. afromontanum (Bullock) 
N. Robson, comb, et stat. nov. 

H. afromontanum Bullock in Kew Bull. 1932: 492 (1932). 

H. annulatum sensu Milne-Redh. in Kew Bull. 8: 435 (1953), 
Fl. Trop. E. Afr., Hypericac.b (1953); Moggi & Pisacchi in 
Webbia 11: 111 (1967) pro parte; Agnew, Upland Kenya 
Wild Fls:l86 (1974); N. Robson in Bamps, Robson & 
Verde, Fl. Trop. E. Afr. Guttif:30 (1978). 



70 



N.K.B. ROBSON 



Stem usually with numerous black (very rarely red) glands, 
densely to sparsely puberulous or rarely glabrous. Leaves 
sometimes with few to numerous laminar black glands, 
puberulous above and densely pubescent beneath or very 
rarely wholly glabrous. Sepals long-glandular-ciliate (more 
than twice as long as glands), usually with some or all laminar 
glands black. Petals with few distal or numerous scattered 
punctiform laminar black glands, always (?) red-tinged in 
bud. 

Southeastern Ethiopia (Harar), East Africa (eastern Uganda, 
southwestern Kenya, northern Tanzania). 

The Harar population is somewhat intermediate between 
subsp. afromontanum and subsp. intermedium, but is more 
similar to the former than the latter. 



REFERENCES 



Arrigoni, P.V., Corrias, S.D., Nardi, E. & Valsecchi, F. 1973. Nuovo stazioni 

di Hypericum annulatum Moris e Ribes multiflorum Kit. ssp. sandalioticum 

Arrig. in Sardegna. Webhia 28:423-425. 
Milne-Redhead, E. 1953a. Tropical African plants: XXIV, Hypcricaceac. Kew 

Bull. 8:434. 
1953ft. Hypcricaceac. Jn W.B. Turrili & E. Milne-Redhead (Eds), Flora 

of Tropical East Africa. London. 
Plumier, C. 1703. Nova plantarum americanum genera: 51. Paris. 
Robson, N.K.B. 1958. The genus Hypericum in Africa south of the Sahara, 

Madagascar and the Mascarenes. Kew Bull. 12:443-446. 
1968. Guttiferae (Clusiaceae). In T.G. Tutin et al. (Eds), Flora Europaea 

2:261-269. 

1977. Studies in the genus Hypericum L. (Guttiferae). 1. Infragencric 

classification. Bull. Br. Mus. nat. Hist. (Bot.) 5:291-355. 

1981. Studies in l\\c genus Hypericum L. (Guttiferae). 2. Characters of the 

genus. Bull. Br. Mus. nat. Hist. (Bot.) 8:55-226. 

1985. Studies in the genus Hypericum L. (Guttiferae). 3 

Campylosporus to 6a. Umbraculoides. Bull. Br. Mus. nat. 
12:163-325. 

1987. Studies in the genus Hypericum L. (Guttiferae). 7. Section 29. 

Brathys (part 1). Bull. Br. Mus. nat. Hist. (Bot.) 16:1-106. 

1990. Studies in the genus Hypericum L. (Guttiferae). 8. Sections 29. 

Brathys (part 2) and 30. Trigynobrathys. Bull. Br. Mus. nat. Hist. (Bot.) 
20:1-151. 



Sections 1. 
Hist. (Bot) 



Bull. nat. Hist. Mus. Lond. (Bot.) 23(2): 71-177 Issued 25 November 1993 



Generic monograph of the 
Asteraceae-Anthemideae 



KARE BREMER 

Department of Systematic Botany, Uppsala University, Villavdgen 6, S -7 52 36 Uppsala, Sweden 

CHRISTOPHER JOHN HUMPHRIES 

Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD 



CONTENTS 



Introduction 73 

Materials and methods 73 

Descriptive terminology 74 

Taxonomic concepts 74 

Cladistics 74 

Tribal and subtribal divisions 74 

Classification 76 

Subtribal classification 76 

Generic classification 77 

Characters of the Anthemideae 78 

Character scoring 78 

Distribution 83 

Key to genera 84 

Anthemideae Cass 90 

1 . Ursiniinae Bremer & Humphries 91 

1 . Ursinia Gaertner 93 

2. Lasiospermum Lagasca 94 

3. Phymaspermum Less 94 

4. Eumorphia DC 94 

5. Gymnopentzia Benth. in Benth. & Hook, f 95 

6. Hymenolepis Cass 95 

7. Athanasia L 95 

8. Asaemia (Harvey) Harvey ex Benth. in Benth. & Hook, f 96 

2. Cancriniinae Bremer & Humphries 96 

9. Trichanthemis Kegel & Schmalh 97 

10. Ugamia Pavlov 98 

11. Richteria Karelin & Kir 98 

12. AllardiaDtcnt 98 

13. Cancr/n/fl Karelin & Kir 99 

14. Cancriniella Tzvelev 99 

3. Tanacetinae' Bremer & Humphries 99 

15. Tanacetum L 100 

16. Opisthopappus Shih 104 

17. Tanacetopsis (Tzvelev) Kovalevsk 104 

18. Xylanthemum Tzvelev 105 

19. LepidolophaWmkX 105 

20. Hippolytia Pol] 105 

21 . Heliocauta Humphries 106 

4. Gonosperminae Bremer & Humphries 106 

22. LMgoflDC 107 

23. Gonospermum Less 107 

24. Inulanthera Kallersjo 108 

5. Handeliinae Bremer & Humphries 108 

25. Lepidolopsis Polj 109 

26. Polychrysum (Tzvelev) Kovalevsk 109 

27. Pseudohandelia Tzvelev 109 

28. Handelia HeimerX HO 

29. Sclerorhachis (Rech. f.) Rech. f HO 

I The Natural History Museum, 1993 



72 K. BREMER AND C. J. HUMPHRIES 

6. Artemisiinae Less, emend. Bremer & Humphries 110 

30. Brachanthemum DC 113 

31. Dendranthema {DC.) Des Moul 113 

32. Arctanthemum (Tzvelev) Tzvelev 114 

33. Tridactylina (DC.) Sch. Bip 115 

34. /1/an/a Polj 115 

35. Phaeostigma Muld 115 

36. Stilpnolepis H. Kraschen 116 

37. Ajaniopsis Shih 116 

38. Filifolium Kitam 116 

39. Sphaeromeria Nutt 117 

40. Kaschgaria Polj 117 

41. Seriphidium (Besser ex Hooker) Four 117 

42. Crossostephium Less 120 

43. Artemisia L 120 

44. Neopallasia Polj 125 

45. Turaniphytum Polj 125 

46. Mausolea Polj 125 

47. Picrothamnus Nutt 125 

7. Achilleinae Bremer & Humphries 126 

48. SantolinaL 128 

49. Otanthus Hoffsgg & Link 128 

50. Achillea L 128 

5 1 . Anacyclus L 129 

52. Leucocyclus Boiss 130 

53. Mecomischus Cosson ex Benth. in Benth. & Hook 130 

54. Chamaemelum Miller 130 

55. Rhetinolepis Cosson 130 

56. Cladanthus Cass 131 

8. Anthemidinae Dumort. emend. Bremer & Humphries 131 

57. Anthemis L 132 

58. Nananthea DC 134 

9. Chrysantheminae Less, emend. Bremer & Humphries 134 

59. Chrysanthemum L 135 

60. Heteranthemis Schott 135 

61 . Ismelia Cass 135 

62. Argyranthemum Webb ex Schultz-Bip 136 

10. Leucantheminae Bremer & Humphries 136 

63. Lepidophorum Necker ex Cass 139 

64. Nipponanthemum Kitam 139 

65 . Leucanthemella Tzvelev 139 

66. Nivellea Wilcox, Bremer & Humphries 140 

67. Phalacrocarpum (DC.) Willk 140 

68. Leucanthemopsis (Giroux) Heyw 140 

69. Hymenostemma (Kunze) Willk 140 

70. Prolongoa Boiss 141 

71 . Leucanthemum Miller 141 

72. Rhodanthemum (Vogt) Bremer & Humphries 141 

73. Leucoglossum Wilcox, Bremer & Humphries 142 

74. Chlamydophora Ehrenb. ex Less 142 

75. Chrysanthoglossum Wilcox, Bremer & Humphries 143 

76. Glossopappus Kunze 143 

77. Coleostephus Cass 143 

78. Plagius L'Herit . ex DC 143 

1 1 . Thaminophyllinae Bremer & Humphries 144 

79. Osmitopsis Cass 145 

80. Adenanthellum B. Nord 145 

8 1 . Inezia E . Phillips 146 

82. Lidbeckia P.J. Bergius 146 

83. Thaminophyllum Harvey 146 

12. Matricariinae Bremer & Humphries 146 

84. Cymbopappus B. Nord 151 

85 . Pentzia Thunb 151 

86. Marasmodes DC 152 

87. Rennera Merxm 152 

88. Oncosiphon Kallersjo 152 

89. Otospermum Willk 153 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 

90. Heteromera Pomel 153 

91 . Daveaua Willk. ex Mariz 153 

92. Matricaria L 153 

93. Microcephala Pobed 154 

94. Endopappus Schultz-Bip 154 

95 . Myxopappus Kallersjo 155 

96. Foveolina Kallersjo 155 

97. Lonas Adans 155 

98. Tripleurospermum Schultz-Bip 155 

99. Aaronsohnia Warb. & Eig 156 

100. Leucoptera B. Nord 157 

101 . Adenoglossa B . Nord 157 

\02.HilliardiaB. Nord 157 

103. CotulaL 157 

104. Leptinella Cass 159 

105. Soliva Ruiz Lopez & Pavon 159 

106. Schistostephium Less 160 

107. Hippia L 160 

108. Eriocephalus L 160 

Excluded genera 161 

References 161 

Index 165 



73 



Synopsis. The Asteraceae tribe Anthemideae is revised. In all, 12 subtribes, 108 genera and 1741 species are 
recognized; nine tribes are described as new and three, the Anthemidinae, the Artemiisinae and the Chrysanthem- 
ineae are emended in circumscription. Four new genera are described and many have been revised in circumscrip- 
tion. The definitions of the tribe, subtribes and genera are expressed in terms of sister group relations and the most 
robust hypotheses of character distribution utilizing the principles of synapomorphy, parsimony and character 
congruence. Phylogenetic relationships were determined using the tree-building computer program, HENNIG86. 
Cladograms of the subtribes and genera are provided together with a synoptic character analysis of each individual 
clade. A key to all genera is provided, each genus is described, all species currently recognized are listed with a brief 
synonomy and the relevant nomenclature and taxonomic changes are discussed in detail. Summaries of distributions 
are given in tables and within the generic accounts. The account ends with a list of excluded taxa and a taxonomic 
index. 



INTRODUCTION 



MATERIALS AND METHODS 



The Asteraceae tribe Anthemideae is one of the largest tribes 
of the family with 1741 species predominantly distributed in 
Eurasia, North and South Africa, with fewer species in North 
America and Australasia. The circumscription of the tribe is 
new but generally follows that outlined by Bentham in 
Bentham & Hooker (1873fl). 

The principal taxonomic problems within the tribe are 
almost entirely relationships between genera but also circum- 
scription of genera especially within subtribal groups such as 
the Artemiisinae, the Chrysantheminae, and the 'Tanaceti- 
nae'. This revision deals with the systematics and generic 
circumscription of the Anthemideae utilizing modern cladistic 
methods. The nomenclature and descriptive taxonomy have 
been fully revised in line with our own analyses and the 
Hterature scanned up to the present time. In addition to the 
formal taxonomic treatment a complete description of charac- 
ter definitions, character distributions and morphologies has 
been provided. Phytogeography and phylogeny have been 
included to give the correct context for the novel classifica- 
tions and a brief historical review of tribal and subtribal 
groupings is also presented. 



The revision is based on herbarium specimens and living 
collections in botanic gardens. The rationale was to study as 
many of the recognized species as possible during the course 
of the work. Those species which we have been unable to 
locate are indicated with an asterisk (*) in the individual lists 
at the end of each generic account. We have studied speci- 
mens from the following herbaria; {Index Herbariorum; 
abbreviations as in Holmgren et al, 1981) BM, BOL, BRY, 
E, F, GH, K, LE, M, MO, NBG, NY, PRE, S, US. Several 
species have been transferred during the course of the 
revision and new combinations are provided. For some North 
African species new combinations based on names in the 
unpublished thesis of Helen Wilcox are validated here. 

The generic descriptions and the character matrices are 
based on dried herbarium specimens and living specimens 
when these were available. Cross-sections of cypselas were 
made from reconstituted collections, softened in water, 
embedded in paraffin and ceresin wax, cut by microtome and 
stained with safranin and light green. The material examined 
has not been listed but all specimens examined were of 
known provenance. 



74 



K. BREMER AND C. J. HUMPHRIES 



Descriptive terminology 

The descriptions and terms used throughout this work follow 
those in Featherly (1954) and Steam (1966). The terminology 
for outlines and plane shapes adopted is that of the Systemat- 
ics Association Committee for Descriptive Biological Termi- 
nology (1962). Descriptions of variation in corolla 
morphology generally follow those of Jeffrey (1977). 

Taxonomic concepts 

The taxonomic concepts in this revision are based on mor- 
phological studies. In most generic revisions, particularly in 
large families such as the Asteraceae, genera and higher taxa 
are frequently circumscribed on uncritical character analyses. 
This approach is considered by us to be unacceptable since in 
no way does it reflect the principles of cladistics. Generic 
names, as with all taxa of any rank, can only be applied to 
monophyletic groups. To recognize monophyletic groups 
which can be designated as genera (or species, or tribes) the 
concept of resemblance has to be resolved so that hypotheses 
of characters can be used to generate hypotheses about 
groups. Evolutionary novelties or synapomorphies, are those 
characters which diagnose monophyletic groups and have 
been utilized here accordingly (see Humphries & Funk, 
1984). 

Cladistics 

The technique we have used to diagnose monophyletic 
groups is the Wagner algorithm as implemented in the 
computer program Hennig86 (Farris, 1988). Hennig86 is an 
interactive program for cladistic analysis which can obtain 
most parsimonious trees by either exact or heuristic calcula- 
tions. Characters were scored from the raw character matri- 
ces into discrete binary codes. Many of the characters we 
used were simply presence or absence characters and thus all 
characters were discretely coded in either a or 1 state. 
However, a considerable number of the characters were 
polymorphic in their distribution especially in the larger 
genera. For precise details of the conventions adopted refer 
to the section on character coding. The character scores are 
presented in twelve matrices (Tables 4, 6, 8, 10, 12, 13, 15, 
17, 19, 21, 23, and 25; see taxonomic accounts below); eleven 
are for individual subtribes (Anthemidineae is combined with 
the Chrysantheminae) and one for subtribes represented as 
individual taxa (Table 4). For the analysis of subtribes the 
matrix (Table 4) was analysed as presented. For the subtribe 
matrices there are three blocks of scores within each matrix; 
the first represents character distributions at a level higher 
than subtribe, the second is characters varying within the 
subtribe and the third provides a summary of ambiguous 
information for a given subtribe. Cladistic analysis for the 
subtribes is based on the central block of characters in each of 
the data matrices. 

Trees were calculated using a variety of options in Hen- 
nig86. All four commands, and five of the nine tree building 
options available in Hennig86 were utilized depending upon 
the size and complexity of each data matrix. The options were 
as follows; hennig (h), which constructs one tree with one 
pass through the data; mhennig* (mh*) which constructs 
several trees, each by a single pass, adding the terminal taxa 
in several different sequences. Branch swapping was applied 
to each of the initial trees and one tree from each initial one 



was retained in the memory; branch-breaker* (bb*) a similar 
option to mhennig* but all of the obtained equally parsimoni- 
ous trees were retained, as was allowed in the assigned 
memory of 128k in the computer; implicit enumeration (ie*) 
which generated all possible shortest trees. The results guar- 
anteed shortest trees and all equal length trees were retained 
in the available tree space; ie- which found one guaranteed 
shortest tree by implicit enumeration. 

Exact analyses (implicit enumeration) were applied to the 
Achilleinae, Anthemidinae, Cancriniinae, Chrysantheminae, 
Gonosperminae, Handeliinae, Tanacetinae, Thaminophylli- 
nae and the Ursiniinae. The hennig, mhennig* and bb* 
options were applied to the larger groups including the 
Anthemideae tribal analysis and the analyses of the Arte- 
misiinae, Leucantheminae and Matricariinae subtribes. The 
trees were all rooted utilizing the outgroup - a taxon scored 
from a North American group of genera within the 
HeHantheae tribe. All scored characters in each of the 
matrices were given equal weight. Unknown scores were left 
in since Hennig86 has the facility for coping with them. In the 
cases where more than one equally parsimonious tree was 
obtained the nelsen command was activated to produce a 
strict consensus tree (Farris, 1988). However, for the purpose 
of the classification only one of the final trees is figured in the 
text (see Figs 1, 2, 7, 11, 17, 20, 23, 31, 36, 41, 47 and 51; see 
taxonomic text below). Each tree was diagnosed using the 
xsteps command in order to determine the states of the 
hypothetical ancestors on each tree (i.e. the node states) and 
to generate statistics on length and fit. 

Precise details of character changes along each clade are 
provided in the subtribe accounts. Each cladogram is pro- 
vided with a unique letter and number code and the length 
and consistency of each tree is provided in the figure caption. 
The cladograms should be used in conjunction with the 
descriptive accounts of the character changes. 

Outgroup 

Bremer (1987) has put forward evidence for the hypothesis 
that Anthemideae and Heliantheae sensu lato are sister 
groups within a larger clade consisting of the Calenduleae, 
the Senecioneae, the Anthemideae and the Heliantheae. 
Bremer also showed that the Anthemideae and its postulated 
relatives in the Heliantheae share similar ray floret epidermis 
as elucidated in the work of Baag0e (1977). The two tribes 
also have similar habit and foliage with trinerved and dis- 
sected leaves especially in the subtribe Helenieae, and thus 
the North American species of this group were selected as the 
outgroup. It has been pointed out to us by Charles Jeffrey (in 
litt.) that the results published by Palmer et al. (1988) from an 
analysis of chloroplast DNA would suggest that we might 
consider a different outgroup. The consensus tree of Palmer 
et al. using 926 restriction site mutations places Astereae and 
Anthemideae as sister groups and the Eupatorieae as an 
ingroup of the Heliantheae. However, we consider that their 
sample of taxa is so preliminary that for the time being we 
remain with Bremer's (1987) morphological hypothesis. 



TRIBAL AND SUBTRIBAL DIVISIONS 

The first coherent tribal classification of the Asteraceae was 
that of Cassini (1817, 1826) who recognized 19 tribes in the 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



75 



family. The precise systematic position of the Anthemideae 
within the Asteraceae is shown in a diagram of the family 
when Cassini used a map with an oval arrangement of 
abutting circles to express the relationships of the tribes (King 
& Dawson, 1975). The Anthemideae were sandwiched 
between the Inuleae and the Ambrosineae. Cassini expressed 
the opinion that the Anthemideae are most closely related to 
the Heliantheae although in style structure they were very 
similar to the Inuleae, the Senecioneae and the Nassauvi- 
ineae. Cassini (1826) divided the tribe into two major groups: 
Anthemidees - Prototypes (Anthemideae-Archetypae) and 
Anthemidees - Prototypes vraies, based on the prescence or 
absence of receptacular scales respectively. For many genera 
the division is quite workable and many of them can be 
placed in one group or the other. However, the artificiality of 
this character can clearly be demonstrated in several taxa 
especially the genus Anthemis {Ammanthus group) in which 
paleate and non-paleate forms can occur within one species 
(Heywood & Humphries, 1977). 

Lessing (1832) had a quite different treatment from that of 
Cassini, and although his classification was avowedly artifi- 
cial, since it was written with identification in mind, it clearly 
influenced the later major treatments of de Candolle (1837), 
Bentham (1873fl, b), and Hoffmann (1894) (see Heywood & 
Humphries, 1977). Lessing recognized eight tribes and the 
genera of the Anthemideae were included in tribe VII 
Senecionideae subtribes Chrysanthemeae and Artemisieae. 
The Chrysanthemeae were further subdivided into two taxa 
of indefinite rank, the Anthemideae and Chrysanthemeae, 
and the Artemisieae were divided up into six taxa of indefi- 
nite rank. 

De Candolle (1837) recognized Cassini's tribe 
Anthemideae as a subtribe and more or less followed 
Lessing's indefinite ranks which he categorized as divi- 
sions. However, he synonymised the Santolineae with the 
Euanthemideae. Bentham (1873a) recognized the 
Anthemideae as a tribe and divided it into six informal 
taxa. Four of these were common to the Lessing and de 
Candolle treatments but he also recognized the Lidbeck- 
ieae and the Tanaceteae as separate taxa. Of other 19th 
century works, that of Hoffmann (1894) is the only one of 
significance and he simply recognized the Cassini classifica- 
tion, dividing the tribe into two subtribes. 

Baillon (1886) provided an end of century classification of 
the angiosperms in his classic Histoire des plantes. For the 
Asteraceae he recognized eight tribes and four hundred and 
three genera. The sub-series Anthemidees was included in an 
enlarged tribe Helianthees together with the Heleniees, 
Senecionees and the true Euhelianthees. 

Most 20th century contributions have been concerned with 
the classification and delimitation of genera but rather silent 
about suprageneric classification and the relationship of the 
tribe to other tribes. However, Bessey (1915) provided 
perhaps the most novel treatment of the Asteraceae when he 
raised the family to the rank of order and raised fourteen 
tribes to family status. Family 296, the Anthemidaceae was 
sandwiched between families 295 and 297, the Eupatoriaceae 
and the Senecionidaceae respectively. However, most treat- 
ments have simply tended to follow the tribal groupings 
outlined by Bentham (1873a) with no real modifications to 
the status of infra-tribal groups (e.g. Carlquist, 1976; Cron- 
quist, 1955, 1977; Wagenitz, 1976). 

Poljakov (1967) recognized six subtribes in the Anthemid- 
eae in a treatment that to some extent resembled that of 



Bentham but with a more detailed consideration of the 
Artemisiinae. 

One of the most recent attempts at creating subgroups 
within the tribe is that of Reitbrecht (1974) who considers 
that the Anthemideae consists of seven provisional groups 
(Humphries, 1979). These include the Ursinia group, to 
include Ursinia and its allies, the Lasiospermum group to 
include the majority of the African genera, especially 
Lasiospermum and Eriocephalus , the Chrysanthemum group 
for some members of the Tanacetinae, Chrysanthemineae 
and Leucanthemineae as recognized by us, the Matricaria 
group to include the Anthemidineae and some members of 
the Matricariinae, and the Cotula group for various other 
members of our Matricariinae. 

Bremer (1987) undertook the first cladistic analysis of the 
tribes of the Asteraceae. The cladogram he obtained sug- 
gested a number of novel groupings. The Anthemideae were 
clearly placed in the monophyletic subfamily, Asteroideae, 
and appeared as the sister group of the Heliantheae in a 
subgroup consisting of the Calenduleae, the Senecioneae, the 
Anthemideae and the Heliantheae, with the Astereae and the 
Eupatorieae as sister groups. Bremer obtained his result by 
considering that the Anthemideae and the Heliantheae share 
similar ray floret cells (Baag0e, 1977), and similar foliage 
with trinerved and dissected leaves which are interpreted as 
synapomorphies for the two tribes. 

Some of the older groups recognized by the 19th century 
authors have caused problems as a result of different inter- 
pretations of characters considered of importance in classifi- 
cation. One of the most problematic groups within the 
Anthemideae is the Ursinia group, and Ursinia in particular. 
Ursinia resembles the Anthemideae in habit and the Arc- 
totideae in terms of superficial features of the capitulum. The 
interpretation as to its exact position has varied, depending 
mostly on the weight of individual characters rather than any 
form of character analysis. Bentham (1873fl) placed Ursinia 
in the tribe Arctotideae (sensu Norlindh, 1977) mainly 
because of its well developed pappus scales. Cassini (1816) 
followed by Beauverd (1915), Merxmuller (1954), Prassler 
(1967), and Reitbrecht (1974) considered it to be a member 
of Anthemideae. Robinson & Brettell (1973) argued that the 
inclusion in the Anthemideae of an anomalous genus like 
Ursinia with its conspicuous pappus scales, widely ovate 
apical anther appendages, and different pollen morphology 
(exine without columnar structure) stretched the workable 
tribal concept. They proposed a new tribe, Ursinieae, which 
included only Ursinia. However, they were not working 
within a cladistic framework and have used in our opinion 
autapomorphic characters to define their new tribe. Accep- 
tance of their solution does nothing to actually clarify the 
position of the Ursinieae in terms of its sister group relations 
and would render the Anthemideae as paraphyletic if the 
Ursinieae were excluded from it. As described in the generic 
account, we consider the large pappus scales and the shape of 
the apical anther appendage plesiomorphic within the 
Anthemideae, since similar structures occur in the outgroup. 
The pollen was investigated by Stix (1960) and she concluded 
that Ursinia belongs to the Anthemideae. The presence of 
unique furanosesquiterpenes which must be considered as 
apomorphic for Ursinia and other South African genera of 
the subtribe Ursiniineae, in our view diagnoses the Ursinieae 
as a natural group and corroborates its tribal position as a 
member group of the Anthemideae. 

Bentham's informal group, the Cotuleae, has also been 



76 



K. BREMER AND C. J. HUMPHRIES 



scrutinized by a variety of workers in recent years. Bentham 
(1873/?) recognized the 'Cotuleae' mostly by 'the mutual 
possession of characters representing loss or reduction in the 
habit and various parts - involucre, paleae, ray florets, 
pappus, number of corolla teeth and stamens (to four) and 
seed-sterility of disk florets' (Lloyd, 1972fl). It is a classic 
polyphyletic group and all of its constituent members (12 
genera; Table 1) have been analysed in terms of gross 
morphology (Heywood & Humphries, 1977), flower and fruit 
structure (Bruhl & Quinn, 1990, 1991), and pollen morphol- 
ogy (Jarvis, 1976; Bruhl & Quinn, 1991). Most of the taxa of 
the Cotuleae are very small plants occurring in the southern 
hemisphere with structurally reduced parts. The detailed 
morphological analyses have shown that most taxa are mis- 
placed within the Anthemideae and these are marked with 
asterisks in Table 1. Several of these have already been 
transferred to other tribes (see list of excluded genera, 
Heywood & Humphries, 1977). The remainder are all mem- 
bers of our revised group, the Matricariinae, in which Cotula, 
Leptinella, and Soliva are all closely related genera. 

In addition to those studies listed by Heywood & Humphries 
(1977), in recent years there have been several studies of higher 
classification in the Anthemideae, particularly by Swedish work- 
ers within the Museum of Natural History, but these have 
concentrated on taxa selected within the tribe. For example, 
Nordenstam (1976^) reclassified a number of genera masquerad- 
ing under the name of Chrysanthemum in South Africa and 
Kallersjo (1988) examined the patterns of relationship in the 
South African Pentzia group of genera. 



CLASSIFICATION 



Subtribal classification 

In this study an entirely new subtribal classification is pro- 
posed. We have grouped genera into larger monophyletic 
entities, subtribes, rather than by a more traditional method 
of dividing the tribe using particular characters. We are fairly 
confident that we have identified a number of monophyletic 
entities, and produced a number of well supported cla- 
dograms. At this time we find it appropriate to recognize 12 
subtribes, most of which represent generic groups that are 
fairly well understood as monophyletic. Several peculiar 
genera of uncertain position have been more or less provi- 
sionally accommodated in otherwise homogenous subtribes. 

Table 1 The 'Cotuleae' group as recognized by Bruhl & Quinn 
(1990, 1991). 



1 


Abrotanella * 


2 
3 
4 


Centipeda * 
Ceratogyne * 
Cotula 


5 
6 


Dimorphocoma * 
Elachanthus * 


7 
8 
9 


Isoetopsis * 

Leptinella 

Nananthea 


10 
11 


Plagiocheilus * 
Soliva 


12 


Tripleurosperm um 


* ^ 


excluded genera 



We think it is better to present a hypothesis on the relation- 
ships of such genera of uncertain position rather than simply 
Hsting them at the end of the tribe. However, genera 
excluded from the Anthemideae are simply listed at the end 
of the revision. 

The predominantly Asian subtribe 'Tanacetinae' is only a 
provisional taxon in this treatment. It is at best paraphyletic 
and will probably turn out to be polyphyletic in later studies. 
It contains the large genus Tanacetum, a dubious taxon of 
critical importance in future studies of the Anthemideae. In 
our study we have grouped together a number of genera that 
appear to have their sister groups within Tanacetum. Further- 
more, it is possible that several other subtribes also have their 
sister groups within Tanacetum. It is clear to us that the 
subtribe 'Tanacetinae', and most particularly Tanacetum 
itself, are unresolved taxa. 

Ursiniinae, Gonosperminae, Handeliinae, Chrysanthemi- 
nae and Thaminophyllinae are homogeneous and monophyl- 
etic subtribes without inclusion of problematic genera. Some 
generic affinities within these subtribes have been understood 
by earlier authors, but in general they are emended here. For 
example, the Canary Island Lugoa and Gonospermum, a well 
known generic pair, are associated with the South African 
Inulanthera, and the three genera form our subtribe Gono- 
sperminae. 

Ursiniinae is the South African group containing fura- 
nosesquiterpenes (Greger, 1977, and references therein) and 
the precise delimitation of the Ursiniinae was undertaken by 
Kallersjo (1986). 

Handeliinae contains a number of small Asian genera. The 
association of Sclerorhachis with the other genera of the 
Handeliinae is new. The affinities between the other genera 
are described already by Tzvelev (in Komarov, 1961) 
although he did not recognize the subtribe and he placed 
them in widely different positions in his floristic classification. 

Chrysantheminae as we understand it is a small group that 
has been identified by earlier authors (see Humphries, 1976; 
Heywood & Humphries, 1977, and references therein). 

Thaminophyllinae is a South African subtribe that hitherto 
has hardly been considered, although affinities of some 
genera have been discussed by recent authors (Nordenstam, 
1976; Bond, 1980). 

Cancriniinae, Artemisiinae, Achilleinae, Anthemidinae, 
Leucantheminae and Matricariinae are subtribes that we also 
consider monophyletic with the character information used in 
this study. Further investigations may result in changes in 
their delimitation, however, since a number of problematic 
genera are involved and the defining characters are rather 
few in some subtribes. Affinities between several of the 
genera within each of these subtribes have been indicated by 
earlier authors but their precise delimitations here are 
entirely new. 

The Asian Cancriniinae seems homogeneous but may 
eventually become a larger group, as its relatives probably 
occur within the undefined subtribe 'Tanacetinae'. The affini- 
ties between some of the genera of the Cancriniinae have also 
been described by Tzvelev (in Komarov, 1961). 

The mainly Asian and large subtribe Artemisiinae contains 
the Artemisia group of genera, an assemblage that has been 
recognized for some time (e.g. Besser, 1829). We have added 
their plesiomorphic relatives to this group. The relationships 
of some of these genera to Artemisia and its relatives have 
been indicated by Poljakov (1955) and Tzvelev in Komarov 
(1961). 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



77 



The Eurasian/North African subtribe Achilleinae is a 
generic group hitherto unrecognized. Interrelationships 
between a few of the genera have been understood (e.g. 
Achillea, Anacyclus, and Leucocyclus, Humphries, 1979; 
Greger, 1977; Valant-Vetschera, 1981) but the subtribe is a 
new generic grouping. Santolina is a plesiomorphic genus of 
uncertain position. It may have close relatives within the 
'Tanacetinae'. 

In this treatment the Anthemidinae contains only two 
genera: Anthemis and Nananthea, the latter a specialized 
genus of uncertain position, and discussed at greater length in 
the generic account. 

The predominantly Mediterranean Leucantheminae 
includes a large subgroup, the Leucanthemum group, which 
has been largely elucidated by the studies of Wilcox (1977) 
and is considered by us to be a well supported monophyletic 
unit. Leucanthemopsis is generally associated with Leucan- 
themum (Heywood, 1954, 1976) and to it we add Phalacro- 
carpum, Hymenostemma and Prolongoa. There are other 
small genera of more uncertain position, such as Lepidopho- 
rum, Nipponanthemum, Leucanthemella and Nivellea. Pend- 
ing further studies, we have considered it worthwhile to 
include them in the Leucantheminae. 

The large cosmopolitan subtribe, Matricariinae, represents 
one of the more interesting generic groupings presented in 
this study. We consider it monophyletic with reservations for 
a few genera of possible uncertain position such as Rennera 
and Oncosiphon. Within the Matricariinae there is the south- 
ern hemisphere Cotula group, an homogeneous, monophyl- 
etic unit, quite distinct from the generic assemblages 
introduced by Bentham (1873fl) and discussed by more recent 
authors (see e.g. Heywood & Humphries, 1977; Bruhl & 
Quinn, 1990, 1991). We have refrained from recognizing the 
Cotula group as a subtribe, since it would probably make the 
remaining Matricariinae paraphyletic. Related to the Cotula 
group are several predominantly Mediterranean and South 
African genera, the interrelationships of which have hitherto 
been unprescribed. 

Although we feel that the majority of our subtribes are well 
supported and easily distinguished, subtribal interrelation- 
ships have been difficult to assess and the main cladogram 
(Fig. 1) is largely unresolved at the subtribal level. The 
cladogram as presented is one of several equally parsimoni- 
ous hypotheses. We suggest that the Ursiniinae and Cancri- 
niinae are plesiomorphic particularly in terms of the scale-like 
pappus structures as compared to other tribes. The genus 
Ursinia within the subtribe Ursiniinae has always been con- 
sidered to occupy an anomalous position in the Anthemid- 
eae. In our opinion, this is partly due to the presence of 
plesiomorphic features. Some of these, particularly basal 
arrangements of leaves and basal woodiness and the internal 
anatomy of cypselas in some members of the Ursiniinae and 
other relatively basal taxa such as the Cancriniinae, are 
remarkably similar to those found in members of the out- 
group which we interpret as examples of symplesiomorphy. 

Within the remaining group of ten subtribes it is reasonable 
to suggest that the predominantly Mediterranean and South 
African subtribes, such as the Achilleinae, Anthemidinae, 
Chrysantheminae, Leucantheminae, Thaminophyllinae and 
Matricariinae are closely related to each other because of 
shared possession of floral resin canals as compared to the 
mainly Asian 'Tanacetinae', Gonosperminae (African but 
apparently directly related to Tanacetum), Handeliinae and 
Artemisiinae. We admit that such a character is vague in 



circumscription but we offer this classification as a basis for 
further investigation. The Thaminophyllinae and Leucan- 
theminae may be closely related subtribes because of similari- 
ties in foliage characters. 

Generic classification 

At the generic level a number of changes should be noted. 
The status of particular genera as being monophyletic or 
non-monophyletic will be discussed briefly. We have adopted 
a conservative style and undertaken generic redelimitations 
only whenever necessary and wherever possible. Our aim has 
been to reclassify para- or polyphyletic genera and maximize 
the number of monophyletic genera within the Anthemideae. 
In many cases we have been content in discussing the 
non-monophyletic status of existing genera, to leave redelimi- 
tations of ambiguous taxa to future systematists. 

Many of the new and redefined genera have been analysed 
in more detail and considered by authors working in close 
connection with us (Kallersjo, 1986, 1988, 1991; Bremer & 
Kallersjo, 1986; Nordenstam, 1976; Ling, 1980a, b, 1991fl, b; 
Wilcox, 1977). The definitions and dispositions of these new 
genera as determined by these authors include Inulanthera 
(Gonosperminae), Rhodanthemum (Leucantheminae), 
Oncosiphon, Myxopappus, Foveolina and Hilliardia (Matri- 
cariinae) and Seriphidium (Artemisiinae). Redefined genera 
include Phymaspermum, Hymenolepis , Athanasia (Ursinii- 
nae), and Pentzia and Rennera (Matricariinae). 

New genera described within this monograph include Chry- 
santhoglossum, Nivellea, Leucoglossum, and Rhodanthe- 
mum, taxa all within the Leucantheminae. Richteria 
(Cancriniinae) and Ismelia (Chrysantheminae) are two small 
genera originally described during the 19th century, later 
reduced into synonomy and then re-established here. Typi- 
cally misplaced species have been transferred to a variety of 
genera such as Microcephala and Aaronsohnia (Matricarii- 
nae). Also, it should be noted that taxonomic changes 
established here have dramatically improved the delimitation 
and definitions of well-known genera such as Artemisia, 
Chrysanthemum (s.s.), Leucanthemum and Tripleurosper- 
mum. By removing a number of species from these formerly 
non-monophyletic genera and assigning the misplaced species 
to new genera, the streamlined versions of the new taxa are 
now more Hkely to be monophyletic. 

The majority of genera within the Anthemideae are mono- 
phyletic taxa but there are several exceptions which will 
obviously require further taxonomic work, as indicated in 
further detail in the generic accounts. Of the relatively 
smaller taxa, genera without autapomorphies include Eumor- 
phia (Ursiniinae), Richteria (Cancriniinae), Tanacetopsis 
('Tanacetinae'), Gonospermum (Gonosperminae), Sphaer- 
omeria (Artemisiinae), Mecomischus and Chamaemelum 
(Achilleinae), Leucanthemopsis and Coleostephus (Leucan- 
theminae), and Cymbopappus and presumably Matricaria 
(Matricariinae). Future work should reveal whether they are 
monophyletic or non-monophyletic taxa. Amongst the larger 
genera, Achillea is one of similar uncertain stature. 

Tanacetum ('Tanacetinae'), Dendranthema, Ajania and 
Artemisia (Artemisiinae), and Cotula (Matricariinae) are 
shown to be non-monophyletic. Other genera and even 
groups of genera have their sister groups within these taxa 
and eventually they will have to be split into smaller mono- 
phyletic units. In the next few years we envisage further 
changes of considerable magnitude which will necessitate 



78 

taxonomic and nomenclatural rearrangements quite different 
from those traditionally recognized. 



CHARACTERS OF THE ANTHEMIDEAE 

The characters used in the cladogram and for delimitation of 
genera are discussed below (see Table 2). The terms charac- 
ter and apomorphy are used synonymously, since any charac- 
ter is apomorphic at its universal level within the taxonomic 
hierarchy. The corresponding plesiomorphic condition is 
omitted from the character table. In obvious cases, such as 
perennial being considered as plesiomorphic as compared to 
annual, the perennial condition is not mentioned. In other 
characters the corresponding plesiomorphic conditions are 
explained elsewhere. The identification of characters and 
character states was undertaken by outgroup comparison 
with parts of the Asteraceae-Heliantheae. 

Character scoring 

There have been problems in scoring character states. For 
several character states, particular variables are expressed 
only within some species of a particular genus. Furthermore, 
the problem of unknown or inapplicable character states has 
caused conflicts during analysis. As far as this study is 
concerned each character may be scored in one of five 
different ways for a particular genus: 

1: The character is present in all species of the genus. 

a: The character is polymorphic and present in only some 
but not necessarily all species of the genus. We take the line 
that all characters of this type are originally present within a 
genus but secondarily lost in some species. This interpreta- 
tion is based on comparison with the immediate relatives of 
the genus, identified during or after cladistic analysis. If the 
character is present in the sister taxa or close relatives, it is 
also considered originally present within the genus; 'a' stands 
for apomorphic. This method is simply outgroup comparison 
with the ingroup restricted to a particular genus being scored 
for the character. 

0: The character is absent from all species of the genus. 

p: The character is absent from some but not necessarily 
most species of the genus. It is interpreted as originally absent 
within the genus; 'p' stands for plesiomorphic but indepen- 
dently derived in other species. The reason for this interpre- 
tation is similar to 'a' above. If the character is absent in the 
relatives, it is also considered originally absent within the 
genus. 

?: The character is unknown or inapplicable to the genus. 
Characters relating to chemistry, embryology and chromo- 
some number are sporadic and unknown for many genera. 
Other characters are inapplicable to some genera. For 
example, characters of receptacular paleae and ray florets are 
inapplicable in epaleate and non-radiate genera, respectively. 
In the cladograms (Figs 1-12; see taxonomic text below) 
unknown or inapplicable characters are assumed to be 
present or absent following the principle of parsimony. If the 
character is known to be present in some genera of a 
particular group, it is also considered present in all genera of 
that group, and vice versa. 



K. BREMER AND C. J. HUMPHRIES 
Table 2 Characters used in the cladograms. 

1 Plants annual. 

2 Plants shrubby. 

3 Plants spiny. 

4 Plants compact and more or less scaphoid. 

5 Plants with one or few sparsely branched stems arising from a 
woody villous caudex. 

6 Plants rhizomatous with rosulate, spathulate-obovate-linear 
leaves. 

7 Plants with branches in whorls below the first capitula. 

8 Plants basally villous-tomentose with rather thick stems and a 
soft pith. 

9 Plants covered with a dense greyish-white indumentum. 

10 Plants covered with viscid hairs. 

11 Plants with dolabriform hairs. 

12 Plants with stellate hairs. 

13 Plants with interxylary cork. 

14 Leaves opposite. 

15 Leaves variously deeply lobed or divided. 

16 Leaves much pinnatisect with filiform lobes. 

17 Leaves pectinate-pinnatisect with filiform, apically somewhat 
swollen and mucronulate lobes. 

18 Leaves with few, oblong to rounded, apically mucronate lobes. 

19 Leaves large with many rounded lobes. 

20 Leaves spathulate in outline, ternate to ternately pinnate. 

21 Leaves serrate-dentate. 

22 Leaves entire or apically tridentate. 

23 Leaves entire, ericoid. 

24 Leaves rather fleshy, few-lobed or entire. 

25 Leaves rather fleshy, entire, linear. 

26 Leaves closely set, lanceolate to linear. 

27 Leaves vermiform. 

28 Leaves with secretory cavities. 

29 Capitula densely corymbose. 

30 Capitula very small and numerous in a large, dense, 
semiglobose corymb. 

31 Capitula in a long narrow panicle or raceme. 

32 Capitula in glomerules arranged in long spikes. 

33 Capitula on short and nodding peduncles. 

34 Capitula sessile along the stems. 

35 Capitula discoid. 

36 Capitula disciform. 

37 Involucre rather narrowly urceolate. 

38 Involucral bracts in 4-7 rows. 

39 Involucral bracts in 1-2 rows, rather wide. 

40 Involucral bracts in 2 unequal series. 

41 Involucral bracts wide, flabelliform. 

42 Involucral bracts subulate. 

43 Involucral bracts with scarious margins. 

44 Involucral bracts with dark brown margins. 

45 Receptacle paieate. 

46 Receptacle narrowly conical to subulate. 

47 Receptacle hollow. 

48 Receptacle pilose. 

49 Receptacle densely hirsute. 

50 Receptacular paleae pilose. 

51 Floral parts with resin canals. 

52 Ray floret limb white. 

53 Ray floret limb golden yellow. 

54 Ray floret limb bluish violet. 

55 Ray floret limb deeply emarginate. 

56 Ray floret hmb epidermis cells tabular (senecioid or mutisioid 
type). 

57 Ray floret tube sinus extending to the base. 

58 Ray and disc floret tube dorsiventrally flattened. 

59 Ray floret tube and cypsela pilose laterally; ray floret limb 
pilose abaxially. 

60 Ray floret tube confluent with the cypsela. 

61 Ray floret tube persistent on the cypsela. 

62 Outer florets stalked. 

63 Outer female florets in several rows. 

64 Outer female florets subtended by scaphoid bracts. 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



79 



Table 2 cont 

65 Outer female floret corollas 'flask-shaped', tapering above or 
narrowly cylindrical. 

66 Outer female floret corollas without teeth. 

67 Outer female florets without corollas. 

68 Outer female floret style-branches lanceolate, flat, acute. 

69 Corolla gradually expanded, rather thin and funnel-shaped. 

70 Corolla inflated with a hollow space between outer surface and 
inner layer. 

71 Corolla apically contracted. 

72 Disc corolla 4-lobed. 

73 Disc corolla lobes with dorsal appendages. 

74 Corollas with continuous veins extending into the lobes. 

75 Disc corolla lobes with central resin sacs. 

76 Disc corolla zygomorphic with 2 smaller adaxial lobes with 
marginal resin canals extending from the base of the corolla 
and with 3 larger abaxial lobes. 

77 Disc corolla red. 

78 Corolla apically with erect, straight hairs. 

79 Corolla apically with long, reddish hairs. 

80 Corolla apically with stellate hairs. 

81 Corolla cobwebby pilose. 

82 Disc corolla tube thickened in fruit. 

83 Disc corolla tube very thick and brittle. 

84 Disc corolla tube basally saccate at least adaxially. 

85 Disc corolla tube deeply and equally saccate both abaxially and 
adaxially. 

86 Corolla tube basally copiously swollen and spongy, almost 
enclosing the cypsela especially laterally. 

87 Disc corolla tube confluent with the cypsela. 

88 Disc corolla tube pilose. 

89 Corolla tube with long-stalked glands; stalk cells elongated. 

90 Disc corolla tube and cypsela ribs with thick vascular strands. 

91 Central florets of two kinds; outer perfect, inner completely 
sterile with reduced ovaries. 

92 Anthers caudate. 

93 Anthers with triangular-linear-lanceolate apical appendages, of 
rather thick-walled cells. 

94 Anthers with an apical resin sac. 

95 Anthers with endothecial tissue partly or wholly polarized. 

96 Pollen grains with short or without spines. 

97 Pollen grains without spines. 

98 Pollen grains hexa-panto-colporate. 

99 Style slender, parallel-sided at base. 

100 Style immersed in a lobed nectary. 

101 Style persistent and spinescent in fruit. 

102 Style-branches brownish. 

103 Disc floret style-branches long-penicillate. 

104 Disc floret style-branches fused. 

105 Stylopodium large and persistent in fruit. 

106 Central floret ovaries reduced; florets functionally male. 

107 Cypselas terete to weakly angled, or flattened. 

108 Cypselas turbinate. 

109 Cypselas arcuate. 

110 Cypselas ellipsoid, small, c. 1 mm long. 

1 1 1 Cypselas subglobose, with 2-3 very thin lateral-adaxial ribs. 

1 12 Cypselas large, with 3 thick protruding sclerenchymatous ribs, 
somewhat winged in ray cypselas. 

1 13 Cypselas slender and tuberculate with numerous obtuse 
excrescences. 

1 14 Cypselas dorsiventrally flattened. 

1 15 Ray cypselas dorsiventrally flattened with 3 adaxial ribs. 

1 16 Disc cypselas laterally flattened. 

1 17 Cypselas laterally winged. 

1 18 Ray cypselas laterally winged; wings projected to apical teeth. 

1 19 Cypselas with sclerenchymatic lateral wings. 

120 Cypselas heteromorphic; ray cypselas triquetrous, winged; disc 
cypselas terete to prismatic to laterally flattened. 

121 Disc cypselas abaxially and adaxially winged. 

122 Cypsela wings as apical spines. 

123 Cypselas with a mainly abaxial entire or toothed rim. 

124 Cypselas with 10 (8-12) multicellular epicarpic ribs. 

125 Cypsela ribs basally fused into a more or less well developed 
foot callus. 

126 Cypsela ribs protruding, narrow and somewhat wing-like. 



Table 2 cont 

127 Cypselas with 5 mainly adaxially arranged ribs. 

128 Cypselas with 1 abaxial and 2 lateral thick ribs and 2-3 adaxial 
ribs. 

129 Cypselas with 1 adaxial and 2 lateral rather thick ribs. 

130 Cypselas with 2 lateral vascular strands, sometimes also with 1 
adaxial strand. 

131 Cypselas 10-ribbed with costal veins and resin canals. 

132 Cypsela ribs with ellipsoid secretory cavities forming 
longitudinal ducts. 

133 Cypselas with costal resin canals or sacs. 

134 Cypselas with vallecular secretory canals. 

135 Cypselas with vallecular vascular strands. 

136 Cypselas abaxially and apically with 2 distinct, occasionally 1 or 
3-5, resin sacs. 

137 Cypselas with a single resin sac apically in the adaxial rib. 

138 Cypselas with scattered elongated resin sacs. 

139 Cypselas completely covered with rows of myxogenic cells. 

140 Cypselas with myxogenic cells on abaxial surface and on the 
ribs of the adaxial surface. 

141 Cypselas with dense rows of myxogenic cells also on the 
corona. 

142 Cypselas with large myxogenic cells in rounded, scattered 
groups. 

143 Cypselas with myxogenic cells in 2 distinct adaxial-lateral rows. 

144 Cypselas with scattered, ovoid, myxogenic trichomes. 

145 Cypselas papillose. 

146 Cypselas long-papillose, seemingly pubescent. 

147 Cypselas densely pilose; hairs subulate, with a few basal cells 
and one long apical cell. 

148 Cypselas cobwebby pilose. 

149 Cypselas with rather stiff unbranched hairs of 3-8 cells with 
spiral wall thickenings. 

150 Cypselas copiously villous, seemingly covered in 'cotton wool'. 

151 Cypsela wall without carbonized layer. 

152 Cypsela wall several cell layers thick, partially or completely 
sclerified. 

153 Cypsela thick-walled and conspicuously rugose. 

154 Cypselas thin-walled, obovoid to oblanceolate, devoid of ribs. 

155 Cypsela wall very thin, translucent and showing brownish 
black, rounded, very thick-walled testa epidermis cells. 

156 Cypsela wall white and spongy; pericarpic cells isodiametric 
with thin spiral wall thickenings. 

157 Cypsela wall with rod-shaped crystals in small packets. 

158 Cypsela wall with numerous druses in the pericarp. 

159 Cypsela wall with a continuous ring of fibre-like cells. 

160 Cypsela wall with a continuous ring of sclerified isodiametric 
cells. 

161 Pappus of short (not large and obovate or bristle-like) scales, 
an auricle, a corona, or absent. 

162 Pappus of scales or teeth projected from the ribs. 

163 Pappus of separate, mainly abaxial, subulate scales. 

164 Pappus of 5 convolute-contorted scales. 

165 Pappus a scarious, flimsy corona. 

166 Pappus adaxially long. 

167 Pappus a stiff adaxial auricle. 

168 Pappus a large, scarious, adaxial but basally coroniform 
auricle, as long as the corolla or longer. 

169 Pappus a large, scarious, adaxial, flabelliform auricle, as long 
as the corolla or longer. 

170 Pappus scales brownish. 

171 Pappus absent in disc cypselas, but present in ray cypselas. 

172 Pappus absent in ray and disc cypselas. 

173 Testa epidermis cells spirally arranged around the embryo. 

174 Testa epidermis cells thick-walled and dark reddish. 

175 Embryo sac tetrasporic. 

176 Embryo sac disporic. 

177 Chromosome number x=10. 

178 'Irregular' monoterpenes in high concentrations present. 

179 Furanosesquiterpenes present. 

180 Particular thiophene derivatives present. 

181 Amides present. 

182 Flavonol 5-glucosides present. 

183 Dehydrofalcarinone and dehydrofalcarinol present. 

184 Anthocyanin present in root tips. 



80 



K. BREMER AND C. J. HUMPHRIES 



Habit 

The outgroup comprises perennial herbs or half-shrubs, as in 
many Anthemideae. Throughout the tribe there are scattered 
annual genera, groups of genera or odd annual species or 
groups of species within otherwise perennial genera. The 
annual habit (character 1) is independently derived several 
times within the Anthemideae. The same holds for the 
shrubby habit (2), although this condition is less common 
within the tribe. No species of the Anthemideae grows into a 
tree. Other uncommon habit characters absent in the out- 
group include characters 3-8. 

Indumentum 

The indumentum of the frequently pubescent Anthemideae 
commonly consists of unbranched hairs with a few basal stalk 
cells and a long apical cell. The same type is found in the 
outgroup and in many other Asteraceae. Occasionally in 
Anthemideae the hairs form a dense, tomentose to villous, 
greyish-white indumentum (character 9). In several 
Anthemideae dolabriform hairs are present (11) comprised of 
a stalk with few cells and transversely arranged apical cells, so 
that the hairs are T-shaped or Y-shaped. A distinction 
between T-shaped and Y-shaped hairs was made by Napp- 
Zinn &. Eble (1980) but we find it difficult to maintain this 
distinction. Both types may be present, sometimes mixed 
together with intermediates. Dolabriform hairs are absent 
from the outgroup. 

In a few genera stellate hairs are present (12), possibly 
derived from the dolabriform hairs. The stellate hairs also 
have a stalk of a few cells, but a stellate apical cell. The type is 
absent from the outgroup. 

Glands composed of two parallel rows of cells with the 
apical pair enlarged are frequent in the Anthemideae, the 
outgroup and other Asteraceae. The number of cell pairs in 
the stalk varies, and at first sight there seem to be two types: 
one with several cells forming a stalk and one with a basal cell 
pair and an apical enlarged pair forming a sessile gland (cf. 
for example, Nordenstam, 1976). There are intermediates 
with two or more basal cell pairs, and hence we find it 
difficult to separate the two types into separate character 
states. Thus, we have avoided using any of them as a 
character. Occasionally, the viscid glands form a dense cover 
on the plant (10). 

Wood anatomy 

In Anthemideae-Artemisiinae a number of genera are pro- 
vided with interxylary cork (character 13; Holmgren et al., 
1976; Moss, 1940). Many genera have not been investigated. 
As far as we know it does not occur in the outgroup. 

Foliage 

In the majority of the Heliantheae the leaves are opposite. 
The outgroup of the Anthemideae within Heliantheae and 
the majority of Anthemideae have alternate leaves. This 
condition then seems best interpreted as a synapomorphy for 
the Anthemideae and part of Heliantheae, whereas the 
secondarily opposite leaf arrangement (character 14) is a 
character within the Anthemideae. 

Variously dissected leaves are characteristic of most 
Anthemideae and entire leaves are rather rare exceptions. 
Dissected leaves also characterize the outgroup, and appear 



to be a synapomorphy (character 15) comprising the 
Anthemideae and the outgroup. Entire leaves then represent 
a reversal in this character. Entire and variously characteristic 
leaves have been scored as independent characters, particu- 
larly 22-26. The mode of leaf dissection is highly variable and 
difficult to apply at the generic level. Various types may be 
present within individual genera. We have designated only a 
few different types as characters which are absent in the 
outgroup, particularly 16-21. 

In Ursiniinae a number of genera have leaves with secre- 
tory cavities, present in floral structures but not in the leaves 
of many other Anthemideae. Secretory cavities are absent in 
the outgroup. 

Inflorescence 

The capitula (inflorescences) are usually terminal either 
singly at the end of a peduncle or aggregated into a variety of 
corymbose, spike-like or clustered synflorescences. Some- 
times the peduncles are structurally reduced so that the 
capitula are aggregated into central sessile clusters (an aggre- 
gated synflorescence) or, on rare occasions, into a single head 
(a syncephalum). The capitula are solitary or arranged in lax 
corymbs in the outgroup and commonly in the Anthemideae. 
They are also frequently arranged in dense corymbs within 
certain genera (character 29, further modified as 30). Rarely, 
the capitula are sessile (34) or solitary on nodding peduncles 
(33). Within the Artemisiinae the corymbose capitulum 
arrangement is further modified into a long narrow panicle- 
like or raceme-like synflorescence (31) generally with numer- 
ous capitula. The paniculate-like capitulum arrangement is 
variable and apparently contains a number of different types. 
The interpretation of these requires a detailed study, and we 
have only listed one distinct type as a character, the arrange- 
ment of the capitula in glomerules on long spikes (32). 

Floral and sex arrangement 

The array of sex expressions and floral morphology in most 
Anthemideae were surveyed by Heywood & Humphries 
(1977). Radiate, heterogamous capitula with hermaphroditic, 
perfect disc florets and female ray florets are the plesiomor- 
phic condition, being present in the outgroup. Sometimes the 
ray florets are neutral but we found this condition difficult to 
apply as a character at the generic level. 

The reduction of the ray florets resulting in discoid, 
homogamous capitula is common, and has occurred several 
times within genera. This character (35) has often been used 
for delimitation of genera, despite its homoplasious nature. 

Disciform, heterogamous capitula (36) with central, her- 
maphroditic florets and outer, female, non-radiate florets 
occur in several Anthemideae. It is possible to hypothesize 
two interpretations for discoid and disciform florets. Outer 
tubular female florets may either be modified central her- 
maphroditic florets or modified female ray florets. The first 
hypothesis implies that disciform capitula are derived modifi- 
cations of discoid capitula. The second hypothesis implies 
that disciform and discoid capitula may be independently 
derived from radiate capitula but it is possible also that 
discoid capitula are derived from disciform capitula by reduc- 
tion of the outer female florets. For analysis it seemed best to 
consider discoid and disciform capitula as independent non- 
additive characters. After construction of the cladogram, a 
particular interpretation may be considered the most parsi- 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



81 



monious, as in the case of the Artemisiinae where discoid 
capitula are considered derived from disciform capitula. In 
other subtribes discoid capitula are best interpreted as 
derived directly from radiate capitula. It was eventually seen 
that there was no support for the hypothesis that disciform 
capitula are derived from discoid capitula in any group of 
Anthemideae. 

Within Artemisiinae the central florets may have reduced 
fertility. In Neopallasia the central florets are heteromorphic. 
The florets towards the periphery are perfect and the central 
ones completely sterile with reduced ovaries (91). In the 
Dracunculus group of the genus Artemisia, central floret 
ovaries are reduced and the florets are functionally male 
(106). 

Involucre 

Involucral shape is variable from widely campanulate to 
narrowly urceolate. In the outgroup the narrow types are 
absent, so that narrowly urceolate involucres are derived 
characters (37) within the Anthemideae. The involucral 
bracts are arranged in several imbricate series as in the 
outgroup and many other Asteraceae. Rarely there are 4-7 
rows (38), more than in the outgroup, or 1-2 rows (39). In 
one genus (Eriocephalus) the involucral bracts are arranged 
in two distinctly unequal rows without intermediates (40). 

In almost all Anthemideae as well as in the outgroup the 
involucral bracts have scarious margins, otherwise they are 
generally absent in the Heliantheae. The chaffy bract is one 
of the main characters (43) which we used to identify the 
outgroup of the Anthemideae. Sometimes the involucral 
bracts are wide and flabelliform (41), or have dark brown 
margins (44), or they are subulate (42), all characters absent 
in the outgroup. 

Receptacle 

The receptacle is either paleate or epaleate. Within the 
Anthemideae the paleate receptacle is plesiomorphic. For- 
merly the tribe Helenieae was distinguished from 
Heliantheae by having epaleate receptacles rather than being 
paleate as in the Heliantheae s.s. Furthermore, the 
Anthemideae was divided into two subtribes based on recep- 
tacular paleae, the paleate Anthemidinae s.l. and the epale- 
ate Chrysantheminae s.l. The homoplasious nature of this 
character has since long been recognized and the distinction 
of the Helenieae and the two subtribes of Anthemideae has 
now been abandoned (see Bremer, 1987). 

It seems possible that the epaleate receptacle is a character 
for a portion of the Heliantheae (s.l.) and Anthemideae. The 
outgroup of the Anthemideae has epaleate receptacles. 
Within Anthemideae the presence of a paleate receptacle is 
hence considered a character (45) despite the fact that it 
might be under single gene control within certain genera (see 
Mitsuoka & Ehrendorfer, 1972). 

The shape of the receptacle varies from being flat to convex 
to conical and ultimately subulate. The latter conditions are 
absent from the outgroup. Receptacle shape is a potential 
source of more character information but we have found it 
difficult to interpret at this stage. It is used as a character (46) 
for a few genera. Rarely the receptacle is hollow (47), pilose 
(48) or densely hirsute (49), characters absent in the out- 
group. The hairs are scattered over the receptacle and similar 
to hairs on other parts, so that they cannot be hypothesized as 



being modified paleae. Occasionally, the paleae are also 
pilose (50) and thus the two characters cannot be considered 
as homologous (failure of the conjunction test, Patterson, 
1982; Humphries & Funk, 1984). 

Resin canals 

In many genera of the Anthemideae floral parts are fre- 
quently invested with resin canals (character 51). They occur 
in a variety of forms, in the corolla lobes, in style-branches, in 
cypselas, sometimes in anther tips and often also in the 
receptacular paleae and involucral bracts. We admit that this 
generalized character is rather vaguely formulated but offer it 
as one hypothesis for suggesting the interrelationships of the 
various subtribes (see discussion below). Resin canals are 
absent in the outgroup. 

Ray florets 

The ray corolla limb is yellow in the outgroup and mostly 
white (character 52) but sometimes yellow in Anthemideae. 
Glossy golden yellow limbs (53) occur in Leucantheminae 
and bluish violet limbs (54) in one genus of the Cancriniinae 
(Allardia), although other colours do occur including pink, 
orange, and red. Occasionally, the Hmb is deeply emarginate 
(55). Baag0e (1977) distinguished a number of epidermal cell 
types on the ray floret limbs. Within the Anthemideae she 
recognized two groups, with Helianthoid and Senecionoid/ 
Mutisioid epidermal cells (56). The former type occurs in the 
outgroup. The ray floret tube furnishes a number of rather 
rare characters not present in the outgroup. The tube is 
sometimes dorsiventrally flattened (58), laterally pilose (also 
abaxially on the limb; 59), confluent with the cypsela (60), 
and persistent on the cypsela (61). Occasionally the ray floret 
tube sinus extends to the base (57), so that logically the tube 
is absent. 

Outer florets in disciform capitula 

Outer female florets and disciform capitula are not present in 
the outgroup. The characters of the outer female florets listed 
here mostly represent unique specializations within Arte- 
misiinae where the plesiomorphic and disciform genus Ajania 
serves as an outgroup. Within the Artemisiinae the outer 
female florets are occasionally subtended by scaphoid bracts 
(character 64), often their corollas are 'flask-shaped' and 
tapering above to narrowly cylindrical (65), and rarely the 
corollas are without teeth (66) or are totally absent (67). 

In disciform Anthemideae the outer female florets are 
sessile and arranged in one row. Rarely in the Matricariinae 
(Cotula group of genera) they are stalked and arranged in 
several rows (Soliva), both very peculiar conditions within 
the family as a whole. The pluriseriate arrangement is com- 
mon in other tribes, but is most parsimoniously interpreted as 
an independent character within the Anthemideae. 

Disc florets 

The corolla of the disc florets provides several characters. In 
the outgroup and in many Anthemideae the plesiomorphic 
condition is seen: yellow, glabrous and actinomorphic, with a 
more or less distinct, unswollen and non-saccate tube, and a 
5-lobed limb without veins or resin sacs in the lobes. 
Within the Ursiniinae the corolla is often gradually 



82 



K. BREMER AND C. J. HUMPHRIES 



expanded and funnel-shaped without distinct tubes and limbs 
(character 69). In Sclerorhachis the limb is contracted at the 
apex (71). In some genera the corolla is 4-lobed (72). The 
lobes may be provided with dorsal appendages (73), continu- 
ous veins along the margins (74), with central resin sacs (75) 
or rarely (Adenoglossa) with marginal resin canals in two of 
the lobes which are smaller than the other three (76). Rarely 
the corolla is red (77) or provided with various types of hairs 
(78-81,88,89). 

The corolla tube is sometimes thickened in fruit (82), and 
in one extreme case (Oncosiphon) very thick and brittle (83). 
It is sometimes saccate basally, adaxially (84) or both abaxi- 
ally and adaxially (85). In an extreme case (Otanthus) the 
tube is copiously swollen at the base and spongy, and almost 
enclosing the cypsela (86). Rarely, the tube may be confluent 
with the cypsela (87, cf. character 60). Other unusual corolla 
types are those with long-stalked glands as found in the 
Ursiniinae (89), and those with very thick vascular strands 
(90). 

Anthers 

In the outgroup and in most Anthemideae the anthers are not 
caudate at the base. This was formerly considered an impor- 
tant character for defining the tribe but anther tails (character 
92) are present in a few genera. The apical anther appendages 
are obtuse to rounded in the outgroup and in most Anthemid- 
eae. In the Artemisiinae they are triangular-linear-lanceolate 
and composed of rather thick-walled cells (93). Sometimes 
the appendages have an apical resin sac (94). The endothecial 
tissue in the outgroup and many Anthemideae and other 
Asteraceae is 'non-polarized', i.e. with the endothecial cell 
wall thickenings arranged evenly. However, in some 
Anthemideae (and other Asteraceae) the endothecial tissue 
is 'polarized' (character 95), with the thickenings arranged 
apically and basally on the cell (Dormer, 1962). 

Pollen 

Spiny pollen is the plesiomorphic condition of the Anthemid- 
eae occurring also in the outgroup. Within the Artemisiinae 
there is a gradual reduction of the spines (characters 96 and 
97) until a rugose appearance is observed on the exine 
surface. Such a condition is considered a modification associ- 
ated with wind-pollination (Wodehouse, 1938). In Adenan- 
thellum, the pollen is hexa-panto-colpororate (98) rather than 
tricolporate as found generally within the Asteraceae. 

Styles 

The disc floret style-branches are of the common so-called 
Senecionoid type: apically penicillate, truncate with parallel 
stigmatic surfaces. In Prolongoa the apical hairs are rather 
long (character 103), and Phaeostigma is named after its 
brownish style-branches (102). In Mausolea (Artemisiinae) 
the style-branches are not linear as in other disciform 
Anthemideae but lanceolate, flat and acute (68). Sometimes 
the style-branches are fused (104) and the style is undivided, 
a condition associated with functionally male florets. 

The style is somewhat bulbous and often situated on a 
more or less developed stylopodium, as in the outgroup and 
many Asteraceae. Within Artemisiinae the style base may be 
slender and parallel-sided (99). Occasionally the stylopodium 
is large and persistent in fruit (105) and in Heliocauta it is 



modified into a lobed nectary (100). In Soliva the style is 
persistent and spinescent in the mature fruit (101). 

Cypselas 

The mature and developing cypselas provide a fruitful supply 
of characters within the Anthemideae (Bruhl & Quinn, 1990; 
Giroux, 1933; Humphries, 1976, 1977; Khandzhyan, 1983; 
Kneisl, 1981; Kynclova, 1970; Wilcox, 1977). In the outgroup 
the cypselas are sharply angled and provided with a heavily 
carbonized layer. This layer is never present in the 
Anthemideae but it is widespread throughout the 
Heliantheae. The reduction of the carbonized layer can be 
interpreted as a synapomorphy (character 151) for the 
Anthemideae as a whole. The cypselas of Anthemideae can 
also be terete to weakly angled, or flattened (107) and 
modified in various ways. The characters 108-113 relate to 
different cypsela shapes unique to the ingroup. The cypselas 
are sometimes much flattened, dorsiventrally (characters 114 
and 115) or laterally (116), and the flattened cypselas are 
often winged (117-122). 

Cypselas without a pappus are apically rounded or truncate 
as in the outgroup, or sometimes provided with an apical rim. 
It is difficult to say whether this rim is homologous with a 
short coroniform pappus or is an independent outgrowth of 
the cypsela wall. We have avoided coding it as a character. In 
Oncosiphon the cypsela rim is distinctly abaxial (123) and 
seems to be a unique structure. 

The presence, number and arrangement of cypsela ribs is 
very variable. Cypselas with five or four evenly arranged ribs 
are common and may be plesiomorphic within the tribe. The 
cypsela ribs are difficult to interpret since similar ribs are 
absent from the outgroup. Characters 124—129 represent 
specialized rib structures within Anthemideae. The cypsela 
ribs are sometimes furnished with resin canals or sacs 
(131-133, 136, 137), not present in the outgroup. Rarely resin 
sacs are scattered over the cypsela (138). Within the Leucan- 
theminae, the Leucanthemum group is characterized by a 
unique cypsela type with vallecular resin canals (134) of a 
particular flattened type and vallecular vascular strands 
(135), situated in the valleys between the ribs, not inside the 
ribs (costal) as in other Anthemideae (Briquet, 1916; Briquet 
& Cavillier, 1916; Giroux, 1933). 

Frequently the cypselas are furnished with myxogenic cells 
within the Anthemideae but the character never occurs in the 
outgroup. However, it is difficult to apply the presence of 
myxogenic cells as a simple presence or absence character as 
it is homoplasious within the tribe as a whole. The myxogenic 
cells are commonly situated along the ribs and this appears to 
be the plesiomorphic arrangement. Other particular distribu- 
tions of the myxogenic cells may serve as characters (139-141 , 
143). Characters 142 and 144 represent unique myxogenic cell 
shapes, quite different from the common type which consists 
of transversely compressed cells in elongated rows. 

The cypselas of the Anthemideae are generally smooth and 
glabrous or sometimes covered with scattered glands. Gla- 
brous and sparsely to rather densely hirsute cypselas are 
found in the outgroup. Papillose cypselas (145) and cypselas 
with other types of indumentum not present in the outgroup 
are those represented by characters 146-150. 

The cypsela wall is sometimes several cell layers thick and 
partially or completely sclerified (152). In other cases it is 
very thin (characters 154 and 155). The intermediate condi- 
tion is common and present in the outgroup. The anatomy of 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 

the cypsela wall is a potential source of more character 
information but detailed investigations of the appropriate 
genera are necessary. The investigations by Reitbrecht (1974) 
need to be expanded and performed in more detail (see Bruhl 
& Quinn, 1990 on the 'Cotuleae'). Characters 156, 159 and 
160 represent unique specializations within the Anthemideae 
(Kallersjo, 1986, 1988). The cells of the cypsela wall usually 
contain isodiametric druses as in many Asteraceae. Rarely 
they are numerous (158) and in Prolongoa they are replaced 
by rod-shaped crystals in small packets (157). 

Pappus 

In the Anthemideae the cypselas are generally without a 
pappus or provided with a coroniform or scaly pappus. True 
bristles, which are never flattened and occur in other tribes, 
are never present. Such bristles are absent also from the 
outgroup, where the pappus, if present, consists of large 
whitish scales or numerous whitish, long, flat bristles (more 
properly bristle-like linear scales). Similar pappus types are 
present in Ursiniinae {Ursinia) and Cancriniinae and we 
assume that the presence of large or long-whitish pappus 
scales are plesiomorphic within the tribe. The five convolute- 
contorted pappus scales (character 164) of Ursinia seem to be 
unique for that genus, although the large size and the whitish 
colour appear to be plesiomorphic. A reduced or lost pappus 
is represented by characters 161, 171 and 172 within the 
Anthemideae. The coroniform pappus is either derived from 
fusion of the individual pappus scales or is an independent 
character unique to the Anthemideae. If we adopt the latter 
hypothesis, logically the coroniform pappus seems to be a 
plesiomorphic condition within the Anthemideae, excluding 
Ursiniinae and Cancriniinae, whereas specialized corona 
types and auricles are apomorphic, particularly as seen with 
characters 162, 163 and 165-170. 



83 

number of x=7, 8, and Artemisia a series based on x=5, 6, 7, 
8 and 9. The highest values have been recorded in Leptinella 
with 2n=312 (Lloyd, 1972a). Most series and most variants 
are aneuploid or polyploid series within genera and variation 
is of Httle use for classification at the generic level. However, 
it appears that the base number of x=10 (character 177) as 
opposed to x=9 is plesiomorphic to a number of genera in the 
Thaminophyllinae and Matricariinae (Cotula group). 

Chemistry 

Chemical characters are difficult to utilize because they are 
mostly collected on a sporadic basis for one or two species 
within a genus. Furthermore, it is rare to find phytochemical 
papers that report absence traits as well as presence of a 
chemical. However, the Anthemideae are well-known for 
accumulating herbal and insecticidal chemicals and relative to 
many angiosperms have been richly studied (Greger, 1975, 
1977; Bohlmann et al., 1973). For systematic purposes avail- 
able chemical evidence suggests that sesquiterpene lactones 
(character 179), flavonoids (182), particularly C-glycosyl fla- 
vones (Valant-Vetschera, 1981, 1982, 1985) and polyacety- 
lenes (183) are of most systematic value (Greger, 1977; 
Tetenyi, 1986). 

Presence of a chemical character has been scored at generic 
rank even though it is acknowledged that investigation has 
been sporadic, particularly with the larger genera, such as 
Artemisia. The scores thus represent hypothetical presence 
for all the species. Different compounds within the same class 
of compounds have been combined into a single score for 
polyacetylenes (183) and furanosesquiterpenes (179) when 
the class compound is considered apomorphic for a particular 
group. Red coloured root tips, due to the presence of 
anthocyanin (184) has been recorded in Leucanthemum 
(Favarger, 1966). 



Testa 

Testa epidermal types were investigated by Kneisl (1981), but 
the results are difficult to interpret. The testa epidermis cells 
are normally well developed with sinuose cells. Sometimes 
the testa collapses during maturation of the fruit. Rarely the 
testa epidermis cells are uniquely specialized, much elon- 
gated and spirally arranged around the embryo (character 
173) or thick-walled and dark reddish (174). 

Embryology 

The Anthemideae have been fairly thoroughly investigated 
compared to many other tribes, by Harling (1950, 1951, 
1960). Tetrasporic embryo sacs (character 175) unambigu- 
ously characterize a number of genera, and dispone embryo 
sacs (176) are reported as unique within Argyranthemum 
(Borgen, 1972). Harling also described development of the 
embryo sacs in detail but the variation is difficult to formulate 
into characters. 

Chromosome numbers 

The Anthemideae have been investigated based on the 
surveys of Federov (1969), Moore (1972, 1973, 1977), and 
Goldblatt (1980, 1981, 1984). X=9 is the base number for the 
tribe. Most genera have either a diploid number of 2n=18 or 
2n=36. Cancrinia has a base number of x=7, Ursinia a base 



DISTRIBUTION 



The approximate total range of the Anthemideae is summa- 
rized in Table 3 and the general distribution of subtribes and 
genera given in Tables 3, 5, 7, 9, 11, 14, 16, 18, 20, 22 and 24; 
see taxonomic text below. The Anthemideae tribe has a 
worldwide distribution but with main concentrations of taxa 
in Central Asia, the Mediterranean region and South Africa. 
Some members of the subtribes Ursiniinae, Artemisiinae, 
Chrysantheminae, Leucantheminae, Anthemidinae, and 
Matricariinae are pernicious weeds, such as some of the 
Ursinia species introduced to Australia and New Zealand, 
and species of Chrysanthemum, Anthemis, Artemisia, Achil- 
lea, Leucanthemum, Tripleurospermum and Matricaria as 
more widespread weeds in both the northern and southern 
hemispheres. However, most taxa have discrete ranges and 
very obvious areas of endemism. The Thaminophyllinae, for 
example, occur only in South Africa. Similarly most Ursinii- 
nae are almost entirely restricted to South Africa, although 
one species of Ursinia occurs in Ethiopia and another species 
of Lasiospermum is found in the Sinai region of Egypt. 

The Gonosperminae is a small subtribe of three genera and 
15 species which displays a presumably relictual distribution 
pattern. Gonospermum and Lugoa are endemic to the 
Canary Islands but have a curious disjunct distribution pat- 
tern with their sister genus, Inulanthera, which has ten species 



84 



K. BREMER AND C. J. HUMPHRIES 



Table 3 Summary of geography and natural distribution of the Anthemideae. 





N.Am. 


Eur- 
Asia 


C.&E. 

Asia 


SW 

Asia 


S.Eur 


N.Afr 


S.Afr 


Austr. 
N.Zeal. 


S.Am 


Number of genera 


8 


25 


38 


13 


28 


38 


29 


3 


4 


Ursiniinae 












2 


8 






Cancriniinae 






6 


1 












Tanacetinae 


1 


2 


6 


2 


1 


1 








Gonosperminae 
Handeliinae 




5 


4 






2 


1 






Artemisiinae 


4 


4 


15 


4 


2 


1 


1 




1 


Achilleinae 




5 


2 


2 


5 


7 








Anthemidinae 




1 




1 


1 


1 








Chrysantheminae 
Leucantheminae 




1 
3 


2 


1 


2 
12 


4 
11 








Thaminophylhnae 
Matricariinae 


3 


4 


3 


2 


5 


10 


5 
14 


3 


3 



in southern Africa, mainly in Natal, but with one species also 
occurring in Madagascar. 

The majority of the remainder of the tribe occur in the 
northern hemisphere. For example, the Chrysantheminae are 
most prominent in North Africa and Macaronesia, although 
the two well-known species of Chrysanthemum, C. coro- 
narium and C. segetum are widespread throughout the north- 
ern hemisphere. Indeed, C. coronarium is cultivated widely 
as a salad vegetable in China. By contrast, the Leucanthemi- 
nae are found in Eurasia and North Africa, and are particu- 
larly concentrated in the Mediterranean region. The 
Achilleinae is more of a Eurasian group with some of the 
more distinctive taxa endemic to North Africa, southern 
Europe, the Mediterranean, and South-West Asia, although 
there are some outlying taxa occuring in North America. The 
'Tanacetinae' are well-represented in Eurasia and especially 
in central Asia but there are some interesting endemics which 
occur also in North America and North Africa. The Cancri- 
niinae and Handeliinae are comprised of eleven genera and 
thirty-four species restricted entirely to the central steppes of 
Asia. The Artemisiinae and Matricariinae are the most 
widespread subtribes. The Artemisiinae which comprise 
about a third of the tribe, with 18 genera and more than six 
hundred species currently recognized, have a worldwide 
distribution but occur mainly in the northern hemisphere and 
especially central and eastern Asia. The Matricariinae, com- 
prised of 25 genera and about 250 species, also has a 
worldwide distribution but most genera occur in the Mediter- 
ranean region and South Africa. 



KEY TO GENERA 

1 Receptacle distinctly paleate with paleae subtending florets (in 
Anthemis rarely with the basal part of the conical receptacle 

epaleate) 2 

Receptacle completely epaleate (though sometimes pilose or 
hirsute) or capitula few-flowered and presence of paleae 
unclear 41 

2 Capitula radiate; rays present 3 

Capitula discoid or disciform ; rays absent 22 

3 Rays yellow, rarely abaxially reddish 4 

Rays white or rarely pink to reddish on both sides 11 

4 Capitula sessile 56. Cladanthus 



Capitula pedunculate 



5 Pappus of 5 (rarely 8-10) large obovate scales and sometimes 5 
additional subulate scales (pappus rarely absent in Ursinia 
trifida, a South African shrub with linear, entire or apically 

few-lobed leaves) 1 . Ursinia 

Pappus a shallow corona, an auricle or rarely of few small 
scales, or absent 6 

6 Disc cypselas 5-ribbed, without pappus; ray cypselas flat and 
sterile with a few pappus scales; leaves serrate 

63. Lepidophorum 

Cypselas equal, without or with a coroniform or auriculiform 
pappus; leaves various 7 

7 Cypselas obovoid without distinct ribs or wings 8 

Cypselas angled or ribbed or prismatic, often turbinate, some- 
times tuberculate; or cypselas flattened and with 2 lateral ribs or 
wings 9 

8 Leaves few-lobed or entire 53 . Mecomischus 

Leaves pinnatifid to pinnatisect 54. Chamaemelum 

9 Cypselas dorsiventrally flattened and with 2 lateral ribs or broad 

wings 10 

Cypselas various, sometimes dorsiventrally flattened but then 
rhombic in cross-section and not or only narrowly winged 

57. Anthemis 

10 Cypselas with 2 lateral more or less distinct ribs but no 

wings 50. Achillea 

Cypselas with 2 lateral wings 51.Anacyclus 

11 Pappus of 5 (rarely 8-10) large obovate scales and sometimes 5 

additional subulate scales 1 . Ursinia 

Pappus a shallow corona, an auricle, or of small scales, or 
absent 12 

12 Cypselas copiously villous 2. Lasiospermum 

Cypselas glabrous, sometimes glandular or papillose 13 

13 Involucral bracts in 2 unequal series; one outer row of pubes- 
cent to glabrescent bracts and one inner row of generally 

densely villous bracts 108. Eriocephalus 

Involucral bracts imbricate and subequal, not in 2 unequal 
series 14 

14 Shrubs with opposite leaves (leaves alternate in Eumorphia 
davyi, a South African shrub with linear, closely set leaves); 

cypselas with 10 or more ribs 4. Eumorphia 

Herbs or rarely shrubs with alternate leaves (rarely with some 
leaves opposite basally on the stems); cypselas various 15 

15 Suffruticose perennial with large leaves with rounded lobes and 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



85 



corymbose capitula; cypselas 5-ribbed with pappus of small 

scales projected from the ribs 22. Lugoa 

Habit and cypselas various but leaves not large and with 
rounded lobes and pappus if present not of scales projected 
mainly from the ribs 16 

16 Cypselas obovoid, without distinct ribs or wings 17 

Cypselas angled or ribbed or prismatic, often turbinate, some- 
times tuberculate; or cypselas flattened and with 2 lateral ribs or 
wings 18 

17 Leaves few-lobed or entire 53. Mecomischus 

Leaves pinnatifid to pinnatisect 54. Chamaemelum 

18 Cypselas dorsiventrally flattened and with 2 lateral ribs or broad 

wings 19 

Cypselas various, sometimes dorsiventrally flattened but then 
rhombic in cross-section and without 2 lateral ribs and not or 
only narrowly winged 21 

19 Cypselas with 2 lateral more or less distinct ribs but no 

wings 50. Achillea 

Cypselas with 2 lateral wings 20 

20 Leaves vermiform; disc corolla tube deeply and equally saccate 

both adaxially and abaxially 52 . Leucocyclus 

Leaves pinnatisect; disc corolla tube adaxially slightly sac- 
cate 51. Anacyclus 

21 Leaves serrate-dentate, rarely pinnatifid or entire; anthers 

caudate 79. Osmitopsis 

Leaves pinnatisect to variously lobed; anthers not 
caudate 57. Anthemis 

22 Pappus present, of scales, an auricle or a corona 23 

Pappus absent, but cypselas sometimes apically with an obtuse 
rim (rarely with a pseudopappus of bristle-like stalked glands in 
Athanasia) 28 

23 Pappus of 5 (rarely 8-10) large obovate scales and sometimes 5 

additional subulate scales 1 . Ursinia 

Pappus a shallow corona, an auricle or of small scales 24 

24 Capitula solitary or laxly corymbose 57 . Anthemis 

Capitula densely corymbose 25 

25 Glabrous annual herb; cypselas with 1 adaxial and 2 lateral ribs 
and with a single secretory cavity apically in the adaxial 

rib 97. Lonas 

More or less pubescent shrubs; cypselas 5-10-ribbed with or 
without several secretory cavities in the ribs 26 

26 Capitula narrowly oblong-obconical, slender, and few-flowered; 

indumentum of stellate hairs 6. Hymenolepis 

Capitula rather widely urceolate to cyathiform-campanulate; 
indumentum of simple or bifid hairs 27 

27 Cypselas 5-ribbed; anthers not caudate 23. Gonospermum 

Cypselas 8-10-ribbed; anthers caudate 24. Inulanthera 

28 Cypselas copiously villous 2 . Lasiospermum 

Cypselas glabrous , sometimes glandular 29 

29 Involucral bracts in 2 unequal series; one outer row of pubes- 
cent to glabrescent bracts and one inner row of generally 

densely villous bracts 108. Eriocephalus 

Involucral bracts imbricate and subequal, not in 2 unequal 
series ^" 

30 Leaves entire or crenate, covered with a dense greyish-white 
indumentum; corolla tube basally copiously swollen and 
spongy, almost enclosing the cypsela especially laterally 

49. Otanthus 

Leaves various; corolla tube not or somewhat swollen and 
spongy but not enclosing the cypsela 31 



31 Cypselas obovoid without distinct ribs or wings 32 

Cypselas angled or ribbed or prismatic, often turbinate, some- 
times tuberculate; or cypselas flattened and with 2 lateral ribs or 
wings 34 

32 Capitula numerous in a long panicle 41 . Seriphidium 

Capitula solitary or laxly corymbose or few closely 
together 33 

33 Capitula almost sessile along the stems 55. Rhetinolepis 

Capitula pedunculate 54. Chamaemelum 

34 Cypselas dorsiventrally flattened and with 2 lateral ribs or broad 

wings 35 

Cypselas various, sometimes dorsiventrally flattened but then 
rhombic in cross-section and not or only narrowly winged . 37 

35 Cypselas with 2 lateral more or less distinct ribs but no 

wings 36 

Cypselas with 2 lateral wings 51. Anacyclus 

36 Leaves mainly rosulate and capitula solitary; cypselas actually 

4-5-ribbed though with 2 major lateral ribs 21. Heliocauta 

Leaves alternate and capitula often corymbose, rarely solitary; 
cypselas with 2 lateral ribs only 50. Achillea 

37 Shrublets; corolla basally saccate around the cypsela especially 

adaxially 48. Santolina 

Herbs or shrubs ; corolla not saccate basally 38 

38 Shrubs with stellate hairs if present; corolla gradually expanded 

and funnel-shaped 7. Athanasia 

Herbs or half-shrubs with simple or bifid hairs; corolla more or 
less distinctly divided into tube and limb 39 

39 Basally villous perennials with much pinnatisect, mainly basal 

leaves; cypselas 4-5-ribbed 40 

Indumentum various but not mainly basal and villous; annuals 
or perennials ; leaves and cypselas various 57 . Anthemis 

40 Stems few-branched, leafy and with terminal corymbs 

28. Handelia 

Stems loosely branched, almost leaf-less with terminal 
capitula 29. Sclerorhachis 

41 Capitula radiate; rays present 42 

Capitula discoid or disciform ; rays absent 109 

42 Rays yellow, rarely partly white or reddish or abaxially red- 
dish 43 

Rays white, rarely pink to reddish, bluish violet or creamy 
orange, but not yellow 62 

43 Pappus present in ray or disc cypselas or generally in all 
cypselas, of scales, an auricle or a corona (sometimes almost 
absent in Tanacetum but cypselas then apically with an acute 

rim) 44 

Pappus absent but cypselas sometimes apically with an obtuse 
rim (in Argyranthemum sometimes with pappus-like apically 
projected cypsela wings) 54 

44 Cypselas densely pilose; pappus of 4-12 white scales at least half 

as long as the corolla 9. Trichanthemis 

Cypselas glabrous, sometimes glandular; pappus of short scales, 
a corona or an auricle 45 

45 Cypselas dorsiventrally flattened and with 2 lateral broad wings; 
an annual herb with entire, fleshy leaves .... 101. Adenoglossa 
Cypselas various, sometimes dorsiventrally flattened but then 
not or only narrowly winged and often rhombic in cross- 
section 46 

46 Disc corolla 4-lobed; leaves entire 81. Inezia 

Disc corolla 5-lobed; leaves various 47 

47 Cypselas with 1 adaxial and 2 lateral distinct ribs, abaxially with 



86 



K. BREMER AND C. J. HUMPHRIES 



or without 2 weaker ribs; pappus coroniform if present 48 

Cypselas various, generally with 5 or more ribs and not with 3 
major adaxial-lateral ribs; pappus and habit various 49 

48 Leaves pinnatifid-pectinate; cypselas with 3 adaxial-lateral thick 
ribs and 2 abaxial weaker ribs; pappus absent in disc cypselas 

but present in ray cypselas 70. Prolongoa 

Leaves pinnatisect; cypselas with 3 adaxial-lateral acute ribs; 
pappus present in both disc and ray cypselas . 94. Endopappus 

49 Cypselas 8-12-ribbed with dark vallecular secretory canals 

between the pale ribs; annuals 50 

Cypselas angled, prismatic or ribbed, but vallecular secretory 
canals absent; annuals or perennials 53 

50 Pappus a scarious, adaxial, flabelliform auricle, as long as the 

corolla or longer 76. Glossopappus 

Pappus a corona, a short auricle, or absent 51 

51 Leaves lobed, often trifurcate; disc cypselas with a stiff coroni- 
form pappus 75. Chrysanthoglossum 

Leaves serrate-dentate; disc cypselas without or with a scarious, 
coroniform or auriculiform pappus 52 

52 Rays golden yellow; cypselas arcuate with the ribs basally and 
adaxially fused into a more or less distinct callus 

77. Coleostephus 

Rays pale yellow; cypselas ellipsoid without a basal callus 

73. Leucoglossum 

53 Cypselas with myxogenic cells on the ribs; pappus a scarious 
flimsy corona; creeping or caespitose suffruticose perenni- 
als 68. Leucanthetnopsis 

Cypselas without myxogenic cells; habit and pappus vari- 
ous 15. Tanacetum 

54 Ray cypselas triquetrous, winged; disc cypselas laterally flat- 
tened and adaxially and abaxially winged or sometimes terete to 

prismatic 55 

Cypselas equal, oblong or obovoid, without wings, or dorsiven- 
trally flattened and laterally winged 58 

55 Shrubs or shrublets 62. Argyranthemum 

Annual herbs 56 

56 Plant covered with viscid hairs; cypsela wings projected to apical 

spines 60. Heteranthemis 

Plants not viscid; cypsela wings if present not projected to apical 
spines 57 

57 Disc corolla red; disc cypselas laterally flattened and adaxially 

and abaxially winged 61. Ismelia 

Disc corolla yellow; disc cypselas prismatic with a narrow 

adaxial wing or terete and apparently ribbed 

59. Chrysanthemum 

58 Disc corolla 4-lobed; rays not true ray florets but outer disc 
florets with one corolla lobe expanded to a limb; cypselas 

dorsiventrally flattened 103. Cotula 

Disc corolla 5-lobed; rays true ray florets with apically 3-lobed 
limbs; cypselas oblong or obovoid 59 

59 Cypselas distinctly ribbed with dark vallecular secretory canals 

between the pale ribs 71. Leucanthemum 

Cypselas distinctly or faintly ribbed but vallecular secretory 
canals absent 60 

60 Basally much woody shrublets with few-lobed leaves with linear 

lobes 30. Brachanthemum 

Herbs or half-shrubs; leaves generally with rather broad 
lobes 61 

61 An annual herb with few-lobed leaves 33. Tridactylina 

Perennials ; leaves various 31. Dendranthema 

62 Pappus present at least in ray cypselas, of scales (bristle-like in 



Allardia), an auricle or a corona (sometimes almost absent in 
ranacemm but cypselas then apically with an acute rim) ... 63 
Pappus absent but cypselas sometimes apically with an obtuse 
rim (mainly abaxial and somewhat toothed in Oncosiphon\ 
cypselas in Argyranthemum sometimes with pappus-like apically 
projected wings) 88 

63 Pappus of 4-12 linear to obovate-oblong scales at least half as 
long as the corolla, or of many bristle-like scales as long as the 

corolla or longer 64 

Pappus of short scales, an auricle or a corona 66 

64 Pappus of 4-12 scales half to equalling the corolla in length; rays 

white or sometimes pink 65 

Pappus of many bristle-like scales equalling or exceeding the 
corolla in length; rays often bluish-violet, sometimes white or 
pink 12. Allardia 

65 Cypselas densely pilose ; pappus scales white 

9. Trichanthemis 

Cypselas glabrous, glandular; pappus scales brownish 

11. Richteria 

66 Cypselas 8-12-ribbed with dark vallecular secretory canals 

between the pale ribs 67 

Cypselas angled, prismatic or often ribbed, but vallecular 
secretory canals absent 69 

67 Annuals; cypselas c. 1 mm long 73. Leucoglossum 

Perennials; cypselas more than 1 mm long 68 

68 Leaves entire or serrate or pinnatifid, sessile; cypsela ribs 

rounded 71 . Leucanthemum 

Leaves trifid or ternate-pinnatifid and seemingly petiolate; 

cypsela ribs narrow and somewhat wing-like 

72. Rhodanthemum 

69 Cypselas triquetrous with 1 adaxial and 2 lateral more or less 
thick ribs and sometimes with 1-2 abaxial weaker ribs, abaxially 
and apically with 2 distinct (occasionally fused to 1 or 3-5) resin 

sacs 98. Tripleurospermum 

Cypselas various, sometimes triquetrous with 3 adaxial-lateral 
ribs but not with 2 abaxial-apical resin sacs 70 

70 Annual herbs 71 

More or less suffruticose perennials, shrublets or shrubs ... 81 

71 Cypselas laterally pilose; pappus a fimbriate whitish corona 

(disc cypselas) or auricle (ray cypselas) 93. Microcephala 

Cypselas glabrous; sometimes glandular; pappus various ... 72 

72 Ray cypselas dorsiventrally flattened and laterally winged; disc 

cypselas 5-ribbed 91. Daveaua 

Cypselas equal or subequal, sometimes somewhat flattened but 
not winged, variously ribbed 73 

73 Cypselas somewhat dorsiventrally flattened and with 2 lateral 

weak ribs; corolla lobes with central resin sacs 

99. Aaronsohnia 

Cypselas with 3 or more, sometimes adaxial, more or less 
distinct ribs, sometimes slightly flattened; corolla lobes without 
or sometimes with central resin sacs (in Matricaria) 74 

74 Cypselas with 1 abaxial (not adaxial) and 2 lateral thick ribs and 
2-3 adaxial weaker ribs ; pappus an adaxial stiff auricle 

89. Otospermum 

Cypselas with mainly adaxial-lateral ribs or with ribs all around 
the cypsela; pappus of scales, a scarious auricle or a corona . 75 

75 Cypsela wall and pappus white and spongy, abaxially thin 

96. Foveolina 

Cypsela wall not white and spongy, pappus scarious 76 

76 Cypselas with 1 adaxial and 2 lateral distinct ribs, abaxially with 
or without 2 weaker ribs, or cypselas with 5 adaxial-lateral 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 

ribs 77 

Cypsela ribs all around the cypsela, not mainly adaxial- 
lateral 79 

77 Cypselas with 1 adaxial and 2 lateral distinct ribs, abaxially with 
orwithout2 weaker ribs; pappus of scales or a stiff corona . 78 
Cypselas with 5 adaxial-lateral ribs; pappus an auricle or a small 
corona 92. Matricaria 

78 Cypselas with 3 adaxial-lateral rounded ribs and 2 abaxial 
weaker ribs, often with 3-5 resin sacs apically in the ribs; pappus 

of 7-10 obovate scales 90. Heteromera 

Cypselas with 3 adaxial-lateral acute ribs; pappus a stiff 
corona 94. Endopappus 

79 Pappus a scarious, flimsy corona; cypselas with myxogenic cells 
on the ribs; leaves pinnatif id-pectinate, spathulate in out- 
line 69. Hymenostemma 

Pappus coroniform, of short more or less connate scales, or an 
auricle; cypselas with or without myxogenic cells; leaves vari- 
ous 80 

80 Cypselas obconical-turbinate, generally with myxogenic cells; 
slender annuals with several stems from a basal rosette (Anthe- 

mis subgen. Ammanthus) 57. Anthemis 

Cypselas more or less oblong, without myxogenic cells; habit 
various 15. Tanacetum 

81 More or less suffruticose perennials 82 

Woody shrubs or shrublets 84 

82 Cypselas with myxogenic cells on the ribs 83 

Cypselas without myxogenic cells 15 . Tanacetum 

83 Pappus a scarious, flimsy corona; leaves serrate to pinnatifid, 
generally pectinate and spatulate in outline 

68. Leucanthemopsis 

Pappus of separate, mainly abaxial scales; leaves pinnati- 
sect 16. Opisthopappus 

84 Cypselas dorsiventrally flattened and laterally winged 

100. Leucoptera 

Cypselas more or less terete, not winged 85 

85 Leaves obovate and apically serrate 64. Nipponanthemum 

Leaves pinnatisect to variously lobed 86 

86 Pappus an adaxially longer cup, or one large adaxial and one 

smaller abaxial scale ; cypselas with myxogenic cells 

84. Cymbopappus 

Pappus a small corona, of several scales or an adaxial auricle; 
cypselas with or without myxogenic cells 87 

87 Cypselas with myxogenic cells; pappus of several adaxially 

longer scales 18. Xylanthemum 

Cypselas without myxogenic cells; pappus a corona of small 
scales or an adaxial auricle 15. Tanacetum 

88 Ray cypselas triquetrous, winged; disc cypselas laterally flat- 
tened and adaxially and abaxially winged 89 

Cypselas generally equal, sometimes triquetrous or dorsiven- 
trally (not laterally) flattened and laterally (not adaxially and 
abaxially) winged 90 

89 Annual herb 61. Ismelia 

Shrubs or shrublets 62. Argyranthemum 

90 Disc corolla 4-lobed 91 

Disc corolla 5-lobed 97 

91 Leaves entire, lanceolate to linear 83. Thaminophyllum 

Leaves variously lobed or pinnatisect, not entire 92 

92 Disc corolla tube very much swollen and brittle 

88. Oncosiphon 

Disc corolla tube not or only slightly swollen 93 



87 

93 Delicate, somewhat succulent, annual herb; leaves with 
rounded lobes; capitula small, 5 mm or less in diam 

58. Nananthea 

Annuals or perennials; leaves various; capitula generally more 
than 5 mm in diam 94 

94 Leaves with few, oblong to rounded, apically mucronate lobes; 

receptacle pilose 82. Lidbeckia 

Leaves variously pinnatisect ; receptacle glabrous 95 

95 Cypselas with 5 adaxial-lateral ribs, somewhat flattened; annu- 
als 92. Matricaria 

Cypselas flattened and with 2 lateral ribs, or obovoid and with 
2-3 faint ribs; perennials or rarely annuals 96 

96 Shrub ; cypselas subglobose with 2-3 adaxial faint ribs 

102. Hilliardia 

Herbs; cypselas dorsiventrally flattened and with 2 lateral 
ribs 103. Cotula 

97 Leaves opposite 98 

Leaves alternate 99 

98 Shrubs with entire or few-lobed leaves 4. Eumorphia 

Creeping, suffruticose perennials with serrate-pinnatifid 
leaves 67. Phalacrocarpum 

99 Cypselas triquetrous, with 1 adaxial and 2 lateral more or less 
thick ribs and sometimes with 2 abaxial weaker ribs, abaxially 
and apically with 2 distinct (occasionally fused to 1 or 3-5) resin 

sacs 98. Tripleurospermum 

Cypselas various, generally with 5 or more ribs, or with 2 lateral 
ribs only 100 

100 Cypselas somewhat dorsiventrally flattened, with 5 adaxial ribs 

or with 2 lateral ribs only 101 

Cypselas more or less terete with 5 or more ribs all around the 
cypsela 102 

101 Cypselas smooth, with 2 lateral weak ribs .... 99. Aaronsohnia 
Cypselas with 5 adaxial-lateral ribs 92. Matricaria 

102 Cypselas distinctly ribbed with dark vallecular secretory canals 

between the pale ribs 71. Leucanthemum 

Cypselas distinctly or faintly ribbed but vallecular secretory 
canals absent 103 

103 Disc corolla tube confluent with the cypsela; cypsela ribs with 

resin canals; leaves serrate 80. Adenanthellum 

Disc corolla tube not confluent with the cypsela; cypsela ribs 
without resin canals; leaves various 104 

104 Cypselas 10-18-ribbed with ovoid myxogenic trichomes; shrubs 

or half-shrubs with entire or few-lobed leaves 

3. Phymaspermum 

Cypselas generally with less than 10 ribs, with or without 
appressed myxogenic cells but not with ovoid myxogenic tri- 
chomes; habit and leaves various 105 

105 Annual herb with lobed, rather lacerate leaves; cypselas without 

myxogenic cells 66. Nivellea 

Perennials; leaves various; cypselas with or without myxogenic 
cells 106 

106 Basally much woody shrublets with few-lobed leaves with linear 

lobes 30. Brachanthemum 

Herbs or half-shrubs generally with rather broad leaf-lobes or 
with linear or serrate leaves 107 

107 Cypselas oblong, more or less distinctly ribbed, without myxo- 
genic cells 108 

Cypselas obovoid, faintly ribbed, mostly with myxogenic 
cells 31. Dendranthema 

108 Leaves mostly rosulate; ray florets fertile . 32. Arctanthemum 
Leaves alternate ; ray florets sterile 65 . Leucantliemella 



K. BREMER AND C. J. HUMPHRIES 



109 Cypselas triquetrous with 1 adaxial and 2 lateral more or less 
thick ribs and sometimes with 2 abaxial weaker ribs, abaxially 
and apically with 2 distinct (occasionally fused to 1 or 3-5) resin 

sacs 98. Tripleurospermum 

Cypselas various, sometimes triquetrous with 3 adaxial-lateral 
ribs but not with 2 abaxial-apical resin sacs 110 

1 10 Pappus present, of scales (bristle-like in Ugamia), an auricle or 
a corona (rarely almost absent in Tanacetum but cypselas then 

apically with an acute rim) Ill 

Pappus absent but cypselas sometimes apically with an obtuse 
rim (mainly abaxial and somewhat toothed in Oncosiphon; 
rarely with a pseudopappus of bristle-like stalked glands in 
Athanasia) 138 

111 Annual herbs 112 

Perennial herbs or often half-shrubs, shrublets or shrubs . 120 

112 Cypselas conspicuously rugose to tuberculate, without ribs> 
apically with a thick, spreading to revolute pappus-like 

rim 87. Rennera 

Cypselas often ribbed and not rugose to tuberculate, or if rugose 
then also with 3 distinct adaxial-lateral ribs; pappus of scales, a 
corona (but not thick and spreading to revolute) or an 
auricle 113 

113 Cypselas somewhat dorsiventrally flattened and with 2 lateral 

weak ribs ; corolla lobes with central resin sacs 

99. Aaronsohnia 

Cypselas with 3 or more, sometimes adaxial, more or less 
distinct ribs, sometimes slightly flattened; corolla lobes without 
or sometimes with central resin sacs (in Matricaria) 114 

114 Cypselas laterally pilose; pappus a fimbriate whitish 

corona 93. Microcephala 

Cypselas glabrous, sometimes glandular; pappus various . 1 15 

115 Cypsela wall and pappus white and spongy, abaxially thin 

96. Foveolina 

Cypsela wall not white and spongy; pappus scarious 116 

116 Cypselas with 1 adaxial and 2 lateral distinct ribs, abaxially with 
or without 2 weaker ribs, or cypselas with 5 adaxial-lateral 

ribs 117 

Cypsela ribs all around the cypsela, not mainly adaxial- 
lateral 118 

117 Cypselas with 1 adaxial and 2 lateral more or less thick ribs; 

pappus a stiff corona covered with myxogenic cells 

95. Myxopappus 

Cypselas with 5 adaxial-lateral ribs; pappus an auricle or a small 
corona 92. Matricaria 

1 18 Cypselas with dark vallecular secretory canals between the pale 
ribs and with myxogenic cells on the ribs; pappus an adaxial but 
basally coroniform auricle as long as the corolla or longer 

74. Chlamydophora 

Cypselas without vallecular secretory canals, with or without 
myxogenic cells; pappus of small scales, a small corona or 
auricle 119 

119 Cypselas obconical-turbinate, generally with myxogenic cells; 
slender annuals with several stems from a basal rosette (Anthe- 

mis suhgen. Ammanthus) 57. Anthemis 

Cypselas more or less oblong, without myxogenic cells; habit 
various 15. Tanacetum 

120 Pappus of 4-15 linear and bristle-like to obovate-oblong scales 

at least half as long as the corolla 121 

Pappus of short scales, an auricle or a corona 123 

121 Cypselas densely pilose; pappus scales white 122 

Cypselas glabrous or sparsely pilose; pappus scales brownish 

13. Cancrinia 



122 Capitula on short, nodding peduncles 10. Ugamia 

Capitula on elongated, straight peduncles or pedunculoid 
stems 9. Trichanthemis 

123 Cypselas with dark vallecular secretory canals between the pale 
ribs and with myxogenic cells on the ribs; pappus an adaxial 

auricle 78. Plagius 

Cypselas without vallecular secretory canals; pappus of scales, a 
corona or sometimes an auricle 124 

124 Capitula small and numerous in an elongated panicle or a large 
corymb; basally villous and woody perennials with much pinna- 

tisect leaves 1 25 

Capitula and leaves various, sometimes small and rather many 
in a panicle or corymb but plant then not basally villous with 
much pinnatisect leaves; perennial herbs, half-shrubs, shrublets 
or shrubs 1 26 

125 Capitula in an elongated panicle 25. Lepidolopsis 

Capitula in a large corymb 26. Polychrysum 

126 Capitula paniculate; leaves apically few-lobed or entire 

42. Crossostephium 

Capitula solitary or laxly to densely corymbose or closely 
aggregated; leaves various 127 

127 A compact, hirsute, basally woody half-shrub with solitary, 
pecunculate capitula; cypselas distinctly 10-ribbed with a coroni- 
form pappus of short wide scales 14. Cancriniella 

Habit and cypselas various, often perennial herbs or 
shrubs 128 

128 Capitula closely aggregated at the stems; leaves small, cricoid, 
entire or occasionally few-lobed; pappus of 7-10 oblong, adaxi- 

ally longer scales 86. Marasmodes 

Capitula solitary or laxly to densely corymbose; leaves and 



pappus various 



129 



129 Capitula narrowly oblong-obconical, slender and few-flowered; 
pubescent shrubs; indumentum of stellate hairs 

6. Hymenolepis 

Capitula generally urceolate to cyathiform or campanulate; 
habit various; indumentum if present of simple or bifid 
hairs 130 

130 Shrub with corymbose capitula and pinnatisect leaves; cypselas 
8-10-ribbed; pappus of small scales projecting from the cypsela 

ribs 24. Inulanthera 

Pappus of scales not distinctly projecting from the cypsela ribs, a 
corona or an auricle ; habit and cypselas various 131 

131 Pollen smooth; a small densely pubescent, basally woody peren- 
nial with solitary or few capitula on short peduncles {Sphaer- 

omeria compacta) 39. Sphaeromeria 

Pollen spiny; habit and capitula various 132 

132 Shrublets with entire or 3-lobed leaves; capitula rather narrowly 
urceolate with involucral bracts in 5-7 rows; pappus of many 

subulate scales 19. Lepidolopha 

Habit, capitula and pappus various, if shrubby with entire or 
3-lobed leaves (some Pentzia species), then not with involucral 
bracts in 5-7 rows and not with a pappus of subulate 
scales 133 

133 Capitula disciform; outer female florets present; cypselas with- 
out myxogenic cells 15. Tanacetum 

Capitula discoid; all florets hermaphrodite; cypselas with or 
without myxogenic cells 134 

134 Corolla tube generally swollen and with thick vascular strands; 

shrubs with more or less cricoid leaves 85. Pentzia 

Corolla tube not or only slightly swollen and with thin vascular 
strands; habit and leaves various 135 

135 Pappus an adaxial auricle or of several adaxially more devel- 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



89 



oped scales 136 

Pappus a corona or of scales, adaxially not more devel- 
oped 137 

136 Perennial herbs with corymbose capitula 15. Tanacetum 

Shrublets with solitary capitula 18. Xylanthemum 

137 Leaves entire but marginally crenate-serrate; perennial herb 

(Tanacetum balsamita) 15. Tanacetum 

Leaves pinnatisect; habit various 17. Tanacetopsis 

138 Leaves opposite; shrubs with sessile or corymbose 

capitula 139 

Leaves alternate, rarely opposite but then habit or capitula 
different 140 

139 Leaves entire; capitula sessile 8. Asaemia 

Leaves pinnatifid; capitula corymbose 5. Gymnopentzia 

140 Capitula solitary or in lax to dense corymbs, small to medium- 
sized and generally erect; pollen usually spiny, rarely smooth (in 
Ajaniopsis, Stilpnolepis, Filifolium, and Sphaeromeria) ... 141 
Capitula in elongated and paniculate, rarely racemiform or 
spiciform or subglobose inflorescences, often small and numer- 
ous and sometimes pendent; pollen smooth 168 

141 Central floret corolla 4-Iobed (rarely 3-lobed in Com/a) ... 142 
Central floret corolla 5-lobed 150 

142 Cypselas dorsiventrally flattened with 2 lateral more or less 

distinct ribs or wings 143 

Cypselas not or only slightly dorsiventrally flattened without or 
with 3-5 ribs and no wings 146 

143 Shrublets or half-shrubs with corymbose or occasionally solitary 

capitula 106. Schistostephium 

Annual or perennial herbs with solitary capitula, rarely few 
together 144 

144 Capitula sessile; outer female florets without corolla and with 

style persistent and spinescent in fruit 105. Soliva 

Capitula more or less pedunculate; outer female florets with 
corolla; style not persistent 145 

145 Outer female floret corolla inflated with a hollow space between 

the outer and inner layer; central florets female-sterile 

104. Leptinella 

Outer female floret corolla not inflated; central florets generally 
fertile 103. Cotula 

146 Perennial (rarely facultative annual) prostrate herbs, regularly 

rooting at the nodes; central florets female-sterile 

104. Leptinella 

Annual herbs, not regularly rooting at the nodes; central florets 
fertile 147 

147 Disc corolla tube very much swollen and brittle 

88. Oncosiphon 

Disc corolla tube not or only slightly swollen 148 

148 DeUcate, somewhat succulent herb; leaves with rounded lobes; 

capitula small, 5 mm or less in diam 58. Nananthea 

Habit and leaves various; capitula generally more than 5 mm in 
diam 149 

149 Cypsela wall either white and spongy with 1 adaxial and 2 lateral 
weak ribs, abaxially thin, or cypselas almost terete and thin- 
walled all around (additional outer conspicuously rugose cypse- 
las may be present) 96. Foveolina 

Cypsela wall not white and spongy, with 5 adaxial-lateral 
ribs 92. Matricaria 

150 Annualherbs 151 

Perennial herbs or often half-shrubs, shrublets or shrubs . 156 



limb inflated, crateriform; cypselas obovoid-lanceolate, densely 

glandular 36. Stilpnolepis 

Involucral bracts obovate-oblong, only marginally scarious; 
corolla limb more or less distinct but not inflated and crateri- 
form; cypselas various but not obovoid-oblanceolate and 
densely glandular 152 

152 Corolla lobes densely pilose; capitula few together in dense 

corymbs; cypselas obovoid with rows of myxogenic cells 

37. Ajaniopsis 

Corolla lobes glabrous, sometimes glandular; capitula often 
solitary or laxly corymbose; cypselas various 153 

153 Cypselas distinctly rugose to tuberculate, more or less 

5-angled 87.Rennera 

Cypselas smooth or ribbed and not rugose to tuberculate (outer 
conspicuously rugose cypselas in addition to the smooth central 
cypselas sometimes present in Foveo/wfl) 154 

154 Cypselas somewhat dorsiventrally flattened and with 2 lateral 

weak ribs ; corolla lobes with central resin sacs 

99. Aaronsohnia 

Cypselas smooth and obovoid to terete, or with 3-5 adaxial- 
lateral, more or less distinct ribs; corolla lobes without or 
sometimes with central resin sacs (in Matricaria) 155 

155 Cypsela wall either white and spongy with 1 adaxial and 2 lateral 
weak ribs, abaxially thin, or almost terete and thin-walled all 
around (additional outer conspicuously rugose cypselas may be 

present) 96. Foveolina 

Cypsela wall not white and spongy, with 5 adaxial-lateral 
ribs 92. Matricaria 

156 Cypselas dorsiventrally flattened, with or without lateral 

wings 107. Hippia 

Cypselas not dorsiventrally flattened, unwinged 157 

157 Cypselas with dark vallecular secretory canals between the pale 
ribs and with myxogenic cells on the ribs .. 71. Leucanthemum 
Cypselas with or without ribs but vallecular secretory canals 
absent 158 

158 Capitula disciform; outer female florets present 159 

Capitula discoid ; all florets hermaphrodite 1 62 

159 Central floret corollas soon compressed together in a resinous 
mass; cypselas obliquely obovoid; a perennial herb with basal 

fibrous leaf sheaths and filiform leaf lobes 38. Filifolium 

Central floret corollas not compressed in a resinous mass; 
cypselas straight and obovoid to obovate-oblong; habit and 
leaves various 160 

160 Pollen smooth; small perennial herbs or half-shrubs 

39. Sphaeromeria 

Pollen spiny (sometimes with short spines only); habit vari- 
ous 161 

161 Style-branches brownish; corolla lobes erect . 35. Phaeostigma 
Style-branches yellowish; corolla lobes spreading .. 34. Ajania 

162 Cypselas slender and somewhat arcuate, tuberculate with 
numerous obtuse excrescences; a basally villous and woody 
perennial with much pinnatisect leaves and densely corymbose 

capitula 27. Pseudohandelia 

Cypselas obovoid-oblong, straight or somewhat oblique, 
smooth or ribbed but not tuberculate; habit, leaves and capitula 
various 163 

163 Cypsela ribs acute, with secretory cavities (best seen in cross- 
section) ; corolla gradually expanded and funnel-shaped 

7. Athanasia 

Cypsela ribs faint or rounded, without secretory cavities; corolla 
more or less distinctly divided into tube and limb 1 64 



151 Involucral bracts widely obovate and largely scarious; corolla 164 Cypselas 10-18-ribbed 3. Phymaspermum 



90 



K. BREMER AND C. J. HUMPHRIES 



Cypselas with fewer than 10 ribs 165 

165 Anthers tailed; perennial herbs with much pinnatisect, basally 

more or less rosulate leaves 20. Hippolytia 

Anthers not tailed; habit various; leaves entire or rather few- 
lobed 166 

166 Perennial herbs with laxly corymbose capitula . 36. Stilpnolepis 
Shrubs or shrublets with solitary capitula 167 

1 67 Leaves with linear lobes 30 . Brachanthemum 

Leaves cricoid with short lobes 85. Pentzia 

168 Capitula disciform; outer female florets present; involucral 

bracts in 2-5 rows 169 

Capitula discoid; all florets hermaphrodite; involucral bracts in 
4-7 rows, unequal, the outer short and rounded, the inner 
gradually longer and linear 41 . Seriphidium 

169 Corolla lobes pilose 170 

Corolla lobes glabrous, sometimes glandular 173 

170 Cypselas densely pilose 171 

Cypselas glabrous 172 

171 Outer female florets without corolla and with dilated, lan- 
ceolate, flat style-branches; a virgate shrub 46. Mausolea 

Outer female florets with a tubular corolla and linear style- 
branches; a woody shrublet with older branches transformed 
into spines 47. Picrothamnus 

172 Leaves pinnatisect; capitula densely congested in glomerules 
arranged in spikes, or solitary in interrupted partly congested 
spikes; indumentum of simple of bifid hairs 

45. Turaniphytum 

Leaves entire or few-lobed; capitula few in an elongated 
panicle, at the summit fasciculate; indumentum of stellate 
hairs 40. Kaschgaria 

173 Suffruticose perennial with entire or 3-5-lobed leaves; capitula 
30-50-flowered in a pyramidal to elongated panicle {Sphaer- 

omeria ruthiae) 39. Sphaeromeria 

Habit, leaves, and capitula various, usually with fewer than 30 
florets 174 

174 Central florets of two kinds; outer perfect, inner completely 
sterile with reduced ovaries; panicle spiciform; leaves pectinate- 
pinnatisect with filiform, apically somewhat swollen and 

mucronulate lobes 44. Neopallasia 

Central florets all perfect or all female-sterile with reduced 
ovaries ; inflorescence and leaves various 43 . Artemisia 



ANTHEMIDEAE Cass. 

J. Phys. Chim. Hist nat. 88: 192 (1819). Type species: 
Anthemis maritima L. 

Aromatic annual or perennial herbs, subshrubs or shrubs, 
rarely spinescent. Leaves alternate or rarely opposite or 
fasciculate or rosulate, generally variously dissected, pinnati- 
sect, pinnatifid, lobed or serrate-dentate or rarely entire. 
Capitula solitary or corymbose or paniculate, rarely aggre- 
gated, often pedunculate, sometimes sessile, variable in size, 
radiate or disciform and heterogamous or homogamous and 
discoid. Involucral bracts in three or more rows, rarely in two 
rows, imbricate, almost always with scarious margins and 
apex. Receptacle paleate or epaleate, rarely pilose or hirsute. 
Ray florets (in radiate capitula) female, fertile or sterile, or 
neuter; limb white, yellow or rarely blue-violet, pink or 
reddish. Outer female florets (in disciform capitula) in one or 



more rows, tubular, rarely without corollas. Central disc 
florets 5- or 4-lobed, rarely 3-lobed, yellow or rarely whitish 
or red, rarely somewhat zygomorphic, perfect or functionally 
male. Anthers generally obtuse at base, rarely shortly tailed; 
filaments basifixed. Style-branches (in central florets) almost 
always truncate and penicillate, with stigmatic areas in two 
marginal stripes, but sometimes undivided in functionally 
staminate florets. Cypselas variable, homo- or heteromor- 
phic, generally terete to weakly angled or ribbed or flattened, 
sometimes winged, thin- or thick-walled, without a carbon- 
ized layer, often with secretory canals and myxogenic cells. 
Pappus generally of rather few scarious scales, a corona or an 
auricle, rarely of many flat bristle-like scales, often absent. 
Embryo sac monosporic or sometimes tetrasporic, rarely 
bisporic. Chromosome number generally x=9, sometimes 
x=10, rarely x=6, 7, 8, 11 or 18. Irregular monoterpenes 
present in high concentrations. 

Distribution (Table 3). Worldwide but with main concen- 
trations in central Asia, the Mediterranean region and South 
Africa. - 12 subtribes, 108 genera, 1741 spp. 

The interrelationships of the 12 proposed subtribes are uncer- 
tain, but one possible hypothesis is presented in the cla- 
dogram (Fig. 1). It should be noted that a number of equally 
parsimonious solutions are possible, and that the strict con- 
sensus tree of these solutions is totally collapsed. The chosen 
cladogram is offered simply as a suggestion for further 
analysis and test. The characters used to identify the various 
clades, indicated in the cladogram, are listed with comments 
below. 



rj= Outgroup 
[=:AO=i=Al|= 1 Ursiniinae 

2 Cancriniinae 
rj= 3 Tanacetinae 
=A2]= 4 Gonosperminae 
h= 5 Handeliinae 



= 6 Artemisiinae 
rj= 7 Achilleinae 

[7= 8 Anthemidinae 



i=A3^ 



:A4:it= 9 Chrysantheminae 
10 Leucantheminae 



|-^A5^ 11 Thaminophyilinae 
1= 12 Matricariinae 



Fig. 1 Cladogram (of five possible) of the 12 subtribes; produced 
by the mhennig* option in Hennig86. Cladogram length = 39, 
consistency index = 84, retention index = 53. 

Clades and characters - Fig. 1, Tables 2 & 4. 

Clade AO 

15 Leaves variously deeply lobed or divided. Dissected leaves 
are characteristic of most Anthemideae as well as of its 
postulated relatives in the Heliantheae. Some of the 
Anthemideae genera and species have entire leaves, but 
these taxa are clearly related to other taxa with dissected 
leaves. 

43 Involucral bracts with scarious margins. This is characteris- 
tic of almost all Anthemideae as well as of its postulated 
relatives in Heliantheae, but also in some austral and 
grangeoid Astereae. At the generic level. Myxopappus with 
subulate involucral bracts represents a reversal. 

Clade Al - tribe Anthemideae 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



91 



Table 4 Data matrix for the Anthemideae. 1 = presence, = 
absence, ? = missing data or not applicable, p = polymorphic but 
scored as the plesiomorphic condition, a = polymorphic but scored 
as the apomorphic condition. 

Ill 1 1 1 1 11 1 111 

1450574597 4616 1319758857 422746 
532718565944192865132124251101706 



1. Ursiniinae 

2. Cancriniinae 

3. Tanacetinae 

4. Gonosperminae 

5. Handeliinae 

6. Artemisiinae 

7. Achilleinae 

8. Anthemidinae 

9. Chrysantheminae 

10. Leucantheminae 

1 1 . Thaminophyllinae 
12.Matricariinae 



alaal?alll00p0000p00pp00p0pp00000 
alaal?0000aap0000p000p000?0000000 
alaalapOOOpplppppppOppOOOppOOpOOp 
alaal?aOp0001 a lOOpOOOOOOO 70000000 
al?al?p000001001al00pp000?0000000 
alaalapOOOpplOOOppalapOOOOpOOOOOO 
alaalaaOOOOplOOOpppOaaalOOppOpOOO 
alaalapOOOOplOOOOppOpapOalpOOOOOp 
alaal 7000000 lOOOpOOOlaOOl pa llpOOO 
alall7pOOOOplOOOOpaOpapOOOppOaOOp 
alall7pp000010000000ppOOO?OOOalOO 
aaallapOpOOOlOOOppOOpapOOpppOOpaa 



52 Ray floret limb white. Rays in the Anthemideae are mostly 
white and sometimes yellow (rarely pink, reddish or blue- 
violet). In the Heliantheae relatives rays are yellow. 

107 Cypselas terete to weakly angled, or flattened. The fruits 
are very variable in the Anthemideae, but the sharply angled 
type, present in all of its postulated immediate relatives in the 
Heliantheae, is hardly ever present. 

151 Cypsela wall without carbonized layer. A carbonized 
layer in the fruit wall is characteristic of many HeUantheae 
and of all Eupatorieae, including the immediate relatives of 
the Anthemideae, but it is totally absent in this tribe. The 
absence of this feature is thus interpreted as a derived 
condition for the tribe. 

178 'Irregular' monoterpenes in high concentrations present 
(Greger, 1977). 

Clade A2 

161 Pappus of short (not large and obovate or bristle-like) 
scales, an auricle, a corona, or absent. In subtribe Cancrinii- 
nae, which is not a member of this clade, the pappus consists 
of many subulate, bristle-like scales and it looks very anoma- 
lous within the tribe. The same pappus type is found in the 
immediate relatives within Heliantheae. Ursinia in subtribe 
Ursiniinae, also not a member of this clade, has a pappus of 
large and obovate scales, similar to some of the wider pappus 
scales in Cancriniinae and Heliantheae. Judging from other 
characters, Ursinia is related to a number of South African 
genera, all united in subtribe Ursiniinae. In these genera the 
pappus is absent and we consider it secondarily lost. How- 
ever, in one genus, Hymenolepis, a pappus of small scales has 
evolved. In all other Anthemideae, comprising clade A2 and 
excluding Cancriniinae and Ursiniinae, the pappus, if not 
secondarily lost, consists of scales, an auricle or a corona. 

Clade A3 

51 Floral parts with resin canals. Resin canals or sacs are 
frequently present in style-branches, corolla lobes and some- 
times also anther tips in a majority of the Anthemideae. They 
seem to occur mainly in the subtribes of this clade, though 
present also in some genera of other subtribes. 

Clade A4 



152 Cypsela wall several cell layers thick, partially or com- 
pletely sclerified. 

Clade A5 

21 Leaves serrate-dentate. In the two subtribes of this clade 
non-dissected leaves dominate, though there are exceptions 
in several genera. 

1. URSINIINAE Bremer & Humphries, subtrib. 
nov. 

Type species: Ursinia paradoxa Gaertner {U. chrysan- 
themoides (Less.) Harvey). 

Herbae annuae vel perennes vel frutices. Corollae floscu- 
lorum radii cellulis epidermaUbus tabularibus. Antherae cel- 
lulis endothecii polaratis. Pappus squamiformis magnus aut 
parvus vel nullus. Furanosesquiterpena adsunt. 

Annual or perennial herbs or shrubs. Leaves alternate or 
sometimes opposite, variable in shape. Capitula solitary or 
laxly to densely corymbose, pedunculate or rarely sessile 
along the stems, radiate or discoid. Receptacle paleate or 
epaleate. Ray floret limb yellow, white or rarely red; epider- 
mis cells tabular. Disc corolla 5-lobed. Anthers with endoth- 
ecial tissue polarized. Cypselas 5- to many-ribbed, glabrous 
or pubescent, papillose to copiously villous, sometimes with 
myxogenic cells. Pappus of large or small scales, or often 
absent. Furanosesquiterpenes present. 

Distribution (Table 5). Southern Africa, one Ursinia spe- 
cies in Ethiopia and one Lasiospermum species in Egypt 
(Sinai), some Ursinia species introduced in Australia and 
New Zealand. - 8 genera, 114 spp. 

Table 5 General distribution of Ursiniinae and genera. 
x=indigenous, o=introduced. 



S. Eur. N. Afr. S. Afr. 



Austr. 
N. Zeal. 



Ursiniinae 
Ursinia 

Lasiospermum 
Phymaspermum 
Eumorphia 
Gymnopentzia 
Hymenolepis 
Athanasia 
Asaemia 



This subtribe was first identified as a group by Bohlmann and 
collaborators (Greger, 1977), who discovered that a number 
of South African Anthemideae genera contained furanoses- 
quiterpenes rather than the common polyacetylenes. Subse- 
quently, the classification of the group has been revised by 
Kallersjo (1986, 1991), who added a number of micromor- 
phological characters (see cladogram in Fig. 2). Kallersjo 
circumscribed it more precisely, by moving a number of 
Athanasia species (to Inulanthera in the Gonosperminae) and 
by including some chemically unknown genera, now placed in 
Phymaspermum and Hymenolepis, which have now been 
investigated chemically. The tribal position of Ursinia within 
Anthemideae, discussed below, has gained further support. 



92 



K. BREMER AND C. J. HUMPHRIES 



There is one alternative equally parsimonious cladogram to 
that shown here. In the alternative cladogram Phymasper- 
mum and Gymnopentzia are sister groups based on a shared 
loss of receptacular paleae (character 45), whereas in the 
presented cladogram Gymnopentzia and Eumorphia are sis- 
ter groups based on their opposite leaves (character 14). 

Clades and characters - Fig. 2, Tables 2, 6. 



|=Url:i(= 1 Ursinia 

[|= 2 Lasiospermum 
lL=Ur2:! r7=Ur4:jj= 3 Phymaspermuin 

l!=Ur5:rf= 4 Eumorphia 

U= 5 Gymnopentzia 
!=Ur3:| [?= 6 Hymenolepis 



7 Athanasia 

8 Asaemia 



Fig. 2 Cladogram (of two possible) of the Ursiniinae produced by 
the ie option in Hennig86. Cladogram length = 39, consistency 
index = 82, retention index = 79. 

Table 6 Data matrix for the Ursiniinae. 1 = presence, = absence, ? 
= missing data or not applicable, p = polymorphic but scored as 
the plesiomorphic condition, a = polymorphic but scored as the 
apomorphic condition. 

Ill 1111111 111 1 1 11 1 1 

145057 4597566735 222441341567395682231 
532718 56599412302894544562194242096344 

I. Ursinia llaal? llllllOpOOOOOOOOOpOOOOOOOOOOOOOO 

2. Lasiospermum 11111? 11110011 llOOOOOOOpOOOOOOOOOOOOOO 

3.Phymaspermuma\l 11? Oil 1001 1001 1 a 1 1 aOpOOOOOOOOOOOOpO 

4. Eumorphia allll? alllO 01 1001 lal lOaOOOOOOOOOOOOOO 

5. Gymnopentzia 11?11? 0? 110011 0011 1 1 101 1 10000000000000 

6. Hymenolepis al?ll? a?1100100011 10000101 11 110000000 

l.Athanasia al?ll? a?110011001 laOOOOlOl llUallalOOO 

%.Asaemia 01?11? 0?110011001100001 10?111111100111 

nil 11 1 

177675555566666171 240 337 
356600357905678511121997853 

\. Ursinia ???000000000000000pppp00000 

2. Lasiospermum ?00??0000000000000p00000000 
3.P/i>'mfl5permMAn??????000000000000000000000 

A. Eumorphia ?????0000000000000000000000 

5. Gymnopentzia ?????????????????0000000000 

6. Hymenolepis ???000???????????00000p0000 

l.Athanasia ?????0???????????00p00pppp0 

%. Asaemia 111 1111 1111 111 1111 {imOQQOQp 



Clade Url - subtribe Ursiniinae 

45 Receptacle paleate. The distribution of this character is 
difficult to interpret. Receptacular paleae are absent in 
Phymaspermum, Gymnopentzia, Asaemia, a few species of 
Athanasia, and one species of Hymenolepis. This is most 
parsimoniously interpreted as secondary losses within the 
subtribe. 

56 Ray floret limb epidermis cells tabular (senecioid or muti- 
sioid type) (Baag0e, 1977). 

95 Anthers with endothecial tissue partly or wholly polarized. 

179 Furanosesquiterpenes present. 

Ursinia 

159 Cypsela wall with a continuous ring of fibre-like cells. 

164 Pappus of 5 convolute-contorted scales. 



Clade Ur2 

161 Pappus of short (not large and obovate or bristle-like) 
scales, an auricle, a corona, or absent. 

172 Pappus absent in ray and disc cypselas. Hymenolepis has a 
pappus of small scales, most parsimoniously interpreted as 
secondarily evolved within this clade. 

Lasiospermum 

133 Cypselas with costal resin canals or sacs. These structures 
are similar to those occurring in the subtribe Matricariinae, 
for example, but different from those in Hymenolepis, Atha- 
nasia, and Asaemia (character 132). 

150 Cypselas copiously villous, seemingly covered in 'cotton 
wool'. 

Clade Ur3 

2 Plants shrubby. 

28 Leaves with secretory cavities. 

29 Capitula densely corymbose. Asaemia and some species of 
Athanasia and Phymaspermum have solitary capitula. 

Clade Ur4 

124 Cypselas with 10 (8-12) multicellular epicarpic ribs. 
Occasionally there are up to 18 ribs. 

145 Cypselas papillose. 
Phymaspermum 

45 reversed. See clade Url. 

144 Cypselas with scattered, ovoid, myxogenic trichomes. In a 
few species the trichomes are not myxogenic, though similar 
in structure. 

Clade Ur5 

14 Leaves opposite. 

Eumorphia 

There is no autapomorphy for Eumorphia. The matter is 
further discussed under the genus. 

Gymnopentzia 

35 Capitula discoid. 

45 reversed. See clade Url. 

146 Cypselas long-papillose, seemingly pubescent. 
Clade Ur6 

12 Plants with stellate hairs. 

35 Capitula discoid. 

51 Floral parts with resin canals. 

69 Corolla gradually expanded, rather thin and funnel-shaped. 

74 Corollas with continuous veins extending into the lobes. 

132 Cypsela ribs with ellipsoid secretory cavities forming 
longitudinal ducts. 

Hymenolepis 

111 reversed. See clade Ur2. 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 

Clade Ur7 

94 Anthers with an apical resin sac. This character occurs in 
most but not all species of Athanasia. 

152 Cypsela wall several cell layers thick, partially or com- 
pletely sclerified. 

160 Cypsela wall with a continuous ring of sclerified isodiamet- 
ric cells. 

Athanasia 

89 Corolla tube with long-stalked glands; stalk cells elongated. 
Some species of Athanasia have glabrous corolla tubes, 
apparently secondarily. 

126 Cypsela ribs protruding, narrow and somewhat wing-like. 

Asaemia 

14 Leaves opposite. 

15 reversed. Leaves not variously deeply lobed or divided, but 
entire. 

23 Leaves entire, cricoid. 

29 reversed. See clade Ur3. 

34 Capitula sessile along the stems. 

45 reversed. See clade Url. 

114 Cypselas dorsiventrally flattened. 

1. URSINIA Gaertner, Fruct. sem. pi. 2: 462 (1791). 
Type species: U. paradoxa Gaertner {V. 
chrysanthemoides (Less.) Harvey). - Sphenogyne R. 
Br. - Ursiniopsis E. Phillips. 

Annual or perennial herbs or shrublets. Leaves alternate, 
entire or generally variously lobed. Capitula solitary or 
laxly corymbose, pedunculate, radiate or occasionally dis- 
coid. Receptacle paleate; paleae scarious, often enveloping 
florets, sometimes narrow with an apical limb. Ray florets 
generally neuter or occasionally female, sterile or fertile; 
limb yellow, white, or rarely red, dorsally often reddish; 
epidermis cells tabular. Disc corolla shallowly 5-lobed. 
Anthers with a widely ovate apical appendage; endothecial 
tissue polarized. Cypselas slender or obovoid, straight or 
curved, 5-ribbed, with a basal tuft of long hairs or glabrous. 
Pappus of 5 (rarely 8-10) large, convolute-contorted, whit- 
ish scales, or of 5 outer such scales and 5 inner subulate, 
whitish scales, or occasionally absent. Furanosesquiterpe- 
nes present. 

Distribution. S. Africa mainly in the SW Cape, also in 
Namibia, Botswana, and Ethiopia. - 38 spp. 

Ursinia was revised by Prassler (1967). Species without a 
basal tuft of cypsela hairs earlier constituted Ursinia s. s. 
(excluding Sphenogyne; the two genera differed also in other 
characters). Species with glabrous cypselas are considered 
derived by Prassler so we assume that presence of cypsela 
hairs is a diagnostic character for the whole genus, though 
secondarily lost in some species. Another former genus, 
Ursiniopsis, was distinguished simply by female rather than 
neutral rays. It was reduced to synonomy by Prassler. Earlier 
authors, e.g. Bentham (1873a), placed Ursinia in the tribe 



93 

Arctoteae sensu Norlindh (1977) mainly because of its well 
developed pappus scales. Cassini (1816) followed by Beau- 
verd (1915) and Prassler considered it a member of 
Anthemideae. Robinson & Brettell (1973) argue that inclu- 
sion in the Anthemideae of the anomalous genus Ursinia with 
its conspicuous pappus scales, widely ovate apical anther 
appendages, and different pollen (exine without columnar 
structure) would destroy a workable tribal concept. Hence 
they proposed a monotypic new tribe, Ursinieae. The large 
pappus scales and the shape of the apical anther appendage 
may be plesiomorphies within Anthemideae, since similar 
structures occur in the outgroup. The pollen was investigated 
by Stix (1960) and she concluded that Ursinia belongs in 
Anthemideae. The presence of unique furanosesquiterpenes 
in Ursinia and a number of other South African Anthemideae 
corroborates its tribal position. 

Although mainly South African, there is one species {U. 
nana) also known from Ethiopia and U. anthemoides is 
introduced into western Australia. The following list of 
species is taken mainly from Prassler's revision. 

U. abrotanifolia (R. Br.) Sprengel 

U. anethoides (DC.) N. E. Br. 

U. anthemoides (L.) Poiret 

U. brachyloba (Kunze) Bremer & Humphries, comb. nov. 

Basionym: Sphenogyne brachyloba Kunze in Linnaea 20: 

21 (1847). 
U. cakilefolia DC. 
U. caledonica (E. Phillips) Prassler 
U. calenduliflora (DC.) N. E. Br. 

U. chrysanthemoides (Less.) Harvey {U. paradoxa Gaertner) 
U. coronopifolia (Less.) N. E. Br. 
U. dentata (L.) Poiret 
U. discolor (DC.) N. E. Br. 
U. dregeana (DC.) N. E. Br. 
U. eckloniana (Sonder) N. E. Br. 
U. filipes (E. Meyer ex DC.) N. E. Br. 
U. frutescens Dinter 
U. heterodonta (DC.) N. E. Br. 
U. hispida (DC.) N. E. Br. 
U. macropoda (DC.) N. E. Br. 
U. merxmuelleri Prassler 
U. montana DC. 
U. nana DC. 

U. nudicaulis (Thunb.) N. E. Br. 
U. oreogena Schltr ex Prassler 

U. paleacea (L.) Moench {U. crithmoides (P. Bergius) Poiret) 
U. pilifera (P. Bergius) Poiret 
U. pinnata (Thunb.) Prassler 
U. punctata (Thunb.) N. E. Br. 
U. pygmaea DC. 

U. quinquepartita (DC.) N. E. Br. 
U. rigidula (DC.) N. E. Br. 
U. saxatilis N. E. Br. 
U. scariosa (Alton) Poiret 
U. sericea (Thunb.) N. E. Br. 
U. serrata (L. f.) Poiret 
U. speciosa DC. 
U. subflosculosa (DC.) Prassler 
U. tenuifolia (L.) Poiret 
U. trifida (Thunb.) N. E. Br. 



94 



K. BREMER AND C. J. HUMPHRIES 



2. LASIOSPERMUM Lagasca, Gen. sp. pi.: 31 
(1816). Type species: L. pedunculare Lagasca (L. 
erectum (Lam.) Druce). 

Annual or perennial herbs. Leaves alternate, variously pinna- 
tisect. Capitula solitary, pedunculate, radiate or discoid. 
Receptacle flat or convex, paleate; paleae thin and scarious 
with a conspicuous resin canal. Ray florets female, fertile; 
limb very short to long, white or reddish; epidermis cells 
tabular. Disc corolla 5-lobed; lobes sometimes reddish. 
Anthers with endothecial tissue polarized. Cypselas copiously 
villous, with resin canals. Pappus absent. Furanosesquiterpe- 
nes present. 

Distribution. S. Africa in the Cape, Namibia, and Egypt in 
Sinai. - 4 spp. 

Lasiospermum is a well defined genus with copiously villous 
cypselas and the development of the cypsela wool deserves 
detailed investigation. One annual species (L. brachyglos- 
sum) occurs also in Sinai. Similar disjunctions are known also 
from other groups. The first three species in the list are taken 
from Flora capensis (Harvey, 1865). 

L. bipinnatum (Thunb.) Druce (L. radiatum Trevir.) 
L. brachyglossum DC. 

L. erectum (Lam.) Druce (L. pedunculare Lagasca) 
L. poterioides Hutch. - Note: Description in Hutchinson, 
1946. 

3. PHYMASPERMUM Less., Syn. gen. compos.: 253 
(1832). Type species: P. junceum Less. - 
Adenachaena DC. - Brachymeris DC. - locaste E. 
Meyer ex Harvey 

Shrubs or half-shrubs. Leaves alternate, entire or lobed, 
often cricoid with secretory cavities. Capitula solitary or 
corymbose, generally pedunculate, rarely sessile along the 
stems, radiate or discoid. Receptacle flat to conical, epaleate. 
Ray florets female, fertile; limb white; epidermis cells tabu- 
lar. Disc corolla 5-lobed, rarely pubescent with long hairs, 
with a narrow tube and a distinct limb. Cypselas 10-18- 
ribbed, generally minutely papillose especially on the ribs and 
generally with scattered ovoid myxogenic trichomes, with a 
more or less distinct apical rim. Pappus absent. Furanoses- 
quiterpenes present. 

Distribution. S. Africa in the Cape, Orange Free State, 
and Transvaal, Swaziland, Zimbabwe and Namibia. - 18 spp. 

Phymaspermum was formerly diagnosed as South African 
shrubs with epaleate and radiate capitula and papillose and 
glandular cypselas. Only the cypsela character represents a 
useful synapomorphy for the genus. Phymaspermum cypselas 
have a peculiar and unique type of myxogenic trichome, 
which are ovoid or almost subglobose and scattered over the 
cypsela surface. Brachymeris was first described as a mono- 
typic genus and the type species, B. scoparia, is a shrub with 
much reduced leaves, small capitula sessile along the stems, 
and a pubescent corolla. Hutchinson (1917) described four 
new species and transferred one Pentzia species to Brachym- 
eris, mainly because they all, like B. scoparia, possessed 
cypselas without a pappus. Hutchinson's species differ from 
B. scoparia in several characters; they have rather closely set 
long leaves, capitula in corymbs, and glandular but not 
pubescent corollas. Kallersjo (1986), who investigated these 



genera, concluded that B. scoparia as well as Hutchinson's 
species could be transferred to Phymaspermum because of 
their similar cypsela morphology. All Brachymeris species 
have the Phymaspermum type of ovoid cypsela trichomes, 
although they are not always myxogenic. Kallersjo also 
transferred a group of Athanasia species with the same 
cypsela morphology to Phymaspermum. These Athanasia 
species are also in habit similar to P. aciculare for example. 
The species of Phymaspermum s. s. (excluding the former 
Athanasia and Brachymeris species but including Aden- 
achaena and locaste) are taken from Flora capensis (Harvey, 
1865) with three species described later added. The former 
Athanasia and Brachymeris species are best identified using 
the treatments by Hilliard (1977) and Hutchinson (1917), 
respectively. 

P. acerosum (DC.) Kallersjo {Athanasia acerosa (DC.) D. 

Dietr. 
P. aciculare (E. Meyer ex Harvey) Benth. ex B. D. Jackson 

{locaste acicularis E. Meyer ex Harvey) 
P. appressum Bolus - Note: Description in Bolus, 1905. 
P. argenteum Brusse - Note: Description in Brusse, 19896. 
P. athanasioides (S. Moore) Kallersjo (Brachymeris athana- 

sioides (S. Moore) Hutch.) 
P. bolusii (Hutch.) Kallersjo {Brachymeris bolusii Hutch.) 
*P. equisetoides Thell. - Note: Description in Thellung, 1923. 
P. erubescens (Hutch.) Kallersjo (Brachymeris erubescens 

Hutch.) 
P. junceum Less. 

P. leptophyllum (DC.) Benth. ex B. D. Jackson (Aden- 
achaena leptophylla DC.) 
P. montanum (Hutch.) Kallersjo (Brachymeris montana 

Hutch.) 
P. parvifolium (DC.) Benth. ex B. D. Jackson (Adenachaena 

parvifolia DC.) 
P. peglerae (Hutch.) Kallersjo (Brachymeris peglerae Hutch.) 
P. pinnatifidum (Oliver) Kallersjo (Athanasia pinnatifida 

(Oliver) Hilliard) 
P. schroteri Compton - Note: Description in Compton, 1931. 
P. scoparium (DC.) Kallersjo (Brachymeris scoparia DC.) 
P. villosum (Hilliard) Kallersjo (Athanasia villosa Hilliard) 
P. woodii (Thell.) Kallersjo - Athanasia woodii (Thell.) 

Hilliard) 

4. EUMORPHIA DC, Prodr. 6: 2 (1838). Type 
species: E. dregeana DC. 

Shrubs. Leaves opposite, rarely alternate, entire or lobed, 
more or less cricoid, with secretory cavities. Capitula gener- 
ally solitary or corymbose, generally pedunculate, radiate. 
Receptacle flat or shghtly convex, rarely conical, generally 
paleate, sometimes epaleate. Ray florets female, fertile; limb 
white; epidermis cells tabular. Disc corolla 5-lobed, with a 
narrow tube and a distinct limb. Anthers with endothecial 
tissue polarized. Cypselas 10-12-ribbed, rarely up to 
18-ribbed, minutely papillose especially on the ribs, with an 
apical rim. Pappus absent. Furanosesquiterpenes present. 

Distribution. S. Africa in the central Cape, Natal and 
Transvaal, and in Lesotho and Swaziland. - 6 spp. 

Traditionally Eumorphia comprises South African 
Anthemideae with radiate capitula (a plesiomorphy), a pale- 
ate receptacle and cypselas without a pappus, characters 
common to several other genera. E. prostrata has a partly 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



95 



epaleate receptacle. The apical cypsela rim present in Eumor- 
phia occurs also in other genera, e.g. Phymaspermum. 
Eumorphia has opposite leaves, which also characterize 
Gymnopentzia. One species, E. davyi, is aberrant in having 
alternate, closely set leaves and a conical receptacle. It is 
provisionally retained in Eumorphia by Kallersjo (1985). The 
circumscription of this genus obviously needs further consid- 
eration. 

E. corymbosa E. Phillips - Note: Description in Phillips, 

1950. 
E. davyi Bolus - Note: Description in Bolus, 1906. 
E. dregeana DC. - Note: Description in Harvey, 1865. 
E. prostrata Bolus - Note: Description in Hilliard, 1977. 
E. sericea J. M. Wood & M. Evans - Note: Description in 

Hilliard, 1977. 
E. swaziensis Compton - Note: Description in Compton, 

1976. 

5. GYMNOPENTZIA Benth. in Benth. & Hook, f., 
Gen. pi. 2(1): 537 (1873). Type species: G. bifurcata 
Benth. 

A shrub. Leaves opposite, pinnatifid, somewhat cricoid, with 
secretory cavities. Capitula corymbose, discoid. Receptacle 
flat or slightly convex, epaleate. Corolla 5-lobed, with a 
narrow tube and a distinct limb. Anthers with endothecial 
tissue polarized. Cypselas 10-ribbed, densely long-papillose 
and thus seemingly pubescent. Pappus absent. Furanoses- 
quiterpenes present. 

Distribution. S. Africa in the E. Cape, Natal and Trans- 
vaal, and in Lesotho. - Monotypic. 

In related genera, i. e. Eumorphia and Phymaspermum, the 
cypselas are minutely papillose, the epidermis cells being 
more or less projected. This is especially pronounced in 
Gymnopentzia, the papillae often being much longer than 
wide and similar to unicellular hairs. The cypselas are thus 
often described as pubescent. The opposite leaves is another 
distinguishing feature from Phymaspermum and possibly 
synapomorphic with Eumorphia. 

6. HYMENOLEPIS Cass, in Bull. Sci. Soc. philom. 
Paris 1817: 138 (1817). Type species: H. parviflora 
(L.) DC. - Phaeocephalus S. Moore. 

Pubescent shrubs; hairs stellate. Leaves alternate, lobed, 
serrate-dentate or entire, with secretory cavities. Capitula 
slender, few-flowered, corymbose, sometimes rather densely 
aggregated, discoid. Receptacle flat, generally paleate, rarely 
epaleate. Corolla gradually expanded and funnel-shaped, 
5-lobed, with short-stalked glands and with continuous veins 
also in the lobes. Anthers with endothecial tissue polarized. 
Cypselas 5-10-ribbed, with ellipsoid secretory cavities form- 
ing longitudinal ducts. Pappus a corona of fimbriate scales. 
Furanosesquiterpenes present. 

Distribution. S. Africa in the Cape. - 7 spp. 

Harvey's (1865) treatment of Hymenolepis as a separate 
genus rather than as a section of Athanasia was recently 
revived by Kallersjo (1986), since Athanasia including Hyme- 
nolepis is paraphyletic. Athanasia s. s. is more closely related 
to Asaemia than to Hymenolepis, as shown in the cladogram 
(Fig. 2). Hymenolepis differs by its slender, few-flowered 



capitula and scaly pappus. The monotypic Phaeocephalus was 
reduced to a synonym of this genus by Kallersjo. A key to the 
species was provided by Bremer & Kallersjo (1986). 

H. cynopus Bremer & Kallersjo 

H. dentata (DC.) Kallersjo {Athanasia schizolepis Harvey) 

H. gnidioides (S. Moore) Kallersjo {Phaeocephalus gnidio- 

ides S. Moore) 
H. incisa DC. 

H. indivisa (Harvey) Kallersjo 
H. parviflora (L.) DC. 
H. speciosa (Hutch.) Kallersjo 

7. ATHANASIA L., Sp. pi. 2nd ed.: 1180 (1763). 
Type species: A. crithmifolia (L.) L. (Bremer & 
Wijnands, 1982). - Stilpnophyton Less. 

Glabrous or pubescent shrubs; hairs stellate. Leaves alter- 
nate, entire, dentate, or lobed, generally cricoid, with secre- 
tory cavities. Capitula generally corymbose, more rarely 
solitary, discoid. Receptacle flat, generally paleate, rarely 
epaleate. Corolla gradually expanded and funnel-shaped, 
5-lobed, glabrous or with long-stalked glands, with continu- 
ous veins also in the lobes. Anthers often with an apical resin 
sac; endothecial tissue polarized. Cypselas 5-12-ribbed gen- 
erally with protruding and narrow ribs, glabrous or occasion- 
ally glandular, with ellipsoid secretory cavities forming 
longitudinal ducts. Pappus absent or often a pseudopappus of 
stalked glands. Furanosesquiterpenes present. 

Distribution. S. Africa, mainly in the SW Cape, one 
species in Natal {A. grandiceps). - 39 spp. 

Athanasia traditionally embraced all South African, discoid 
Anthemideae with a paleate receptacle. Epaleate species 
were placed in other genera, e. g. Pentzia and Stilpnophyton. 
Athanasia and Stilpnophyton (as well as Asaemia) have 
similar cypsela wall anatomy, with a continuous ring of 
sclerified isodiametric cells. Stilpnophyton was reduced to a 
synonym by Kallersjo (1986). 

Several former species of Athanasia have now been trans- 
ferred by Kallersjo to the genera Hymenolepis, Phymasper- 
mum, and Inulanthera. The remaining part, Athanasia s. s. is 
a homogeneous and monophyletic group. Characteristically, 
the cypsela ribs are narrow and somewhat wing-like, and the 
cypselas are often furnished with a peculiar 'pseudopappus' 
of long-stalked glands. In some species such glands are also 
present on the corolla tube, and in others they are totally 
absent, probably secondarily. In her most recent treatment of 
Athanasia (Kallersjo, 1991) the distinctive characters of 
Asaemia are considered to be autapomorphic and A. minuta 
is considered to be very similar to Athanasia humilis 
Kallersjo. The list of species is based on Kallersjo (1986, 
1991). 

A. adenantha (Harvey) Kallersjo 

A. alba Kallersjo 

A. bremeri Kallersjo 

A. calophylla Kallersjo 

A. capitata (L.) L. 

A. cochlearifolia Kallersjo 

A. crenata (L.) L. 

A. crithmifolia (L.) L. 

A. cuneifolia Lam. 

A. dentata (L.) L. 



96 

A. elsiae Kallersjo 

A. filiformis L. f. 

A. flexuosa Thunb. 

A. grandiceps Milliard & Burtt 

A. hirsuta Thunb. 

A. humilis Kallersjo 

A. imbricata Harvey 

A. inopinata (Hutch.) Kallersjo {Stilpnophyton inopinatum 

Hutch.) 
A. juncea (DC.) D. Dietr. 
A. leptocephala Kallersjo 
A. linifolia Burm. (Stilpnophyton linifolium (L. f.) Less., 

Stilpnophyton longifolium (Thunb.) Less.) 
A. microcephala DC. 
A. microphylla DC. 
A. minuta (L. f.) Kallersjo 
A. oocephala (DC.) Kallersjo {Stilpnophyton oocephalum 

DC.) 
A. pachycephala DC. 
A. pectinata L. f. 
A. pinnata L. f. 
A. pubescens (L.) L. 
A. quinquedentata Thunb. 
A. rugulosa E. Meyer ex DC. 
A. scabra Thunb. 
A. sertulifera DC. 
A. spathulata (DC.) D. Dietr. 
A. tomentosa Thunb. 
A. tr if areata (L.) L. 
A. vestita (Thunb.) Druce 
A. virgata Jacq. 
A. viridis Kallersjo 

8. ASAEMIA (Harvey) Harvey ex Benth. in Benth. & 
Hook, f., Gen. pi 2(1): 433 (1873). Type species: 
A. axillaris (Thunb.) Harvey ex Hoffmann {A. 
minuta (L. f.) Bremer). 

A glabrous, sometimes spiny shrub. Leaves opposite, sheath- 
ing, entire, cricoid, with secretory cavites. Capitula sessile, 
solitary along the branches and on lateral branchlets, discoid. 
Receptacle flat, epaleate. Corolla gradually expanded and 
funnel-shaped, 5-lobed, glandular, with continuous veins also 
in the lobes. Anthers with an apical resin sac; endothecial 
tissue polarized. Cypselas 2-5-angled but generally dorsiven- 
trally flattened with 1 adaxial and 2 lateral ribs, glabrous or 
basally with a few hairs, with few ellipsoid secretory cavities 
forming longitudinal ducts, apically with a smooth or denticu- 
late thickened rim. Pappus absent. Furanosesquiterpenes 
present. 



Distribution. 
Monotypic. 



S. Africa in the Cape and in Namibia. - 



Asaemia minuta is habitually very distinct, a shrub with 
cricoid leaves and small sessile capitula. It is sometimes 
spinescent (ssp. minuta) and sometimes unarmed (ssp. iner- 
mis (E. Phillips) Bremer). Asaemia is related to Athanasia. 
They have similar cypsela wall anatomy with a continuous 
ring of sclerified isodiametric cells (Kallersjo, 1986). Both 
Asaemia and most species oi Athanasia also have anthers with 
apical resin sacs. The genus was revised by Bremer (1983) but 
sunk into Athanasia by Kallersjo (1991). 



K. BREMER AND C. J. HUMPHRIES 

2. CANCRINIINAE Bremer & Humphries, 
subtrib. nov. 

Type species: Cancrinia chrysocephala Karelin & Kir. 

Plantae perennes, herbaceae vel suffruticosae, compactae, 
plusminusve scaphoideae. Bracteae involucri plerumque mar- 
gine atrofuscae. Pappus e squamis vel setis planis pluribus, 
obovatis vel hnearibus, albidis vel brunneis, longitudine 
quam corolla 2-plo brevioribus vel longioribus formatus. 

Compact, more or less scaphoid perennial herbs or half- 
shrubs. Leaves alternate to rosulate, pinnatifid to pinnatisect. 
Capitula solitary, pedunculate, radiate or discoid. Involucral 
bracts generally with dark brown margins. Receptacle epale- 
ate, glabrous or sometimes pilose or hirsute. Ray floret limb 
white, yellow, pinkish or bluish violet. Disc corolla 5-lobed, 
sometimes pilose. Cypselas 5-15-ribbed, glabrous or pilose. 
Pappus of several, obovate to linear, whitish or brownish 
scales or flat bristles, at least half as long as the corolla. 



Distribution (Table 7). 
genera, 26 spp. 



Asia, mainly central part. - 6 



Table 7 General distribution of Cancriniinae and genera. 
x=indigenous, o=introduced. 





C.&E. 


SW 




Asia 


Asia 


Cancriniinae 


X 


X 


Trichanthemis 


X 




Ugamia 


X 




Richteria 


X 


X 


Allardia 


X 




Cancrinia 


X 




Cancriniella 


X 





This subtribe comprises some of the most plesiomorphic 
representatives of the tribe Anthemideae. In habit species of 
Triehanthemis and Richteria, and in pappus structure Ugamia 
and Allardia, are similar to members of the outgroup in tribe 
Heliantheae. Most species are restricted to mountain habitats 
in central Asia. Ugamia, Cancriniella and some Trichanthe- 
mis species, are small, compact, much woody, cushion- 
formed half-shrubs. Cancrinia and Allardia are compact 
perennial herbs. This habit character, together with the dark 
brown-margined involucral bracts, are both considered syna- 
pomorphies for the genera of the subtribe. It is possible that 
groups of species in Tanacetum, subtribe 'Tanacetinae', also 
sharing these features, are related to members of Cancrinii- 
nae rather than 'Tanacetinae', and should be transferred to 
Cancriniinae. 

The cladogram for the genera is only one of several equally 
parsimonious ones, and shown only to display one possible 
hypothesis of generic interrelationships. The strict consensus 
tree for the alternative cladograms is totally collapsed. 

Clades and characters - Fig. 3, Tables 2, 8. 

Clade Cal - subtribe Cancriniinae 

4 Plants compact and more or less scaphoid. Allardia and 
Cancrinia are compact, more herbaceous perennials with 
rather short peduncles, whereas the other genera are much 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



97 



:Ca2JC 



— 14 



9 Trichanthemis 

10 Ugamia 

11 Richteria 

12 Allardia 

13 Cancrinia 



Cancriniella 



Fig. 3. Cladogram (of 21 possible) of the Cancriniinae produced 
by the ie option in Hennig86. Cladogram length = 14, consistency 
index = 85, retention index = 60. 

Table 8 Data matrix for the Cancriniinae. 1 = presence, = absence, 
? = missing data or not applicable, p = polymorphic but scored as 
the plesiomorphic condition, a = polymorphic but scored as the 
apomorphic condition. 



Ill 
145057 
532718 



1 1 1 

44483375346 
447983504981 



11 11 

15775555566666175 
30563567905678511 



9. Trichanthemis WaW? lallaOpOOOOO ????000000000000p 

10. Ugamia al?al? allOOllOOOOO ????????????????0 

n. Richteria alaal? alOOOOOlOOOO ????000000000000p 

U.Allardia alaal? alOOOOOllOOO ????0000000000000 

13. Cancrinia al?al? alOOOOl 10000 ????????????????p 

U. Cancriniella al?al? alOOOOlOOlll ????????????????0 



woody basally, generally with long peduncles. Ugamia has 
short peduncles. 

44 Involucral bracts with dark brown margins. This character 
is absent in some Trichanthemis species. 

Clade Ca2 

147 Cypselas densely pilose; hairs subulate, with a few basal 
cells and one long apical cell. Cancrinia and some species of 
Allardia sometimes have sparsely pilose cypselas. 

Trichanthemis 

49 Receptacle densely hirsute. 

88 Disc corolla tube pilose. Some species of Trichanthemis 
have glabrous corollas. 

Ugamia 

33 Capitula on short and nodding peduncles. 

35 Capitula discoid. Some species of Trichanthemis, Can- 
crinia and Cancriniella are also discoid. 

Clade Ca3 

170 Pappus scales brownish. 

Richteria 

There is no obvious autapomorphy for this genus, and it 
appears undefined compared to Allardia and Cancrinia. 

Allardia 

54 Ray floret limb bluish violet. It is not clear if this character 
occurs in all Allardia species. Some may have whitish rays. 

Cancrinia 

35 Capitula discoid. 

Cancriniella 

35 Capitula discoid. 

39 Involucral bracts in 1-2 rows, rather wide. According to 



Tzvelev (in Komarov, 1961) this character distinguishes Can- 
criniella from Cancrinia. 

48 Receptacle pilose. 

161 Pappus of short (not large and obovate or bristle-like) 
scales, an auricle, a corona, or absent. 



9. TRICHANTHEMIS Regel & Schmalh. in Trudy 
imp. S.-Peterb. hot. Sada 5: 617 (1877). Type 
species: T. karataviensis Regel & Schmalh. - 
Glossanthis Polj . 

Basally much woody half-shrubs with erect annual stems 
basally covered in sheathing leaf bases. Leaves alternate and 
basally rosulate, pinnatisect. Capitula soUtary, pedunculate, 
radiate or discoid. Involucral bracts with or without dark 
brown margins. Receptacle convex, generally densely hir- 
sute, epaleate. Ray florets female, fertile; limb white, pale 
pink, or yellow, many-veined; tube generally pilose. Disc 
corolla 5-lobed, generally pilose; hairs subulate, with a few 
basal cells and one long apical cell. Cypselas 7-10-ribbed, 
densely pilose with the same type of hairs as on the florets, 
sometimes with myxogenic cells. Pappus of 4— 12 large, white, 
linear to oblong scales at least half as long as the corolla. 

Distribution. C. Asia. - 9 spp. 

Trichanthemis has several unusual features such as the hirsute 
receptacle and the pilose corollas and cypselas. The relation- 
ship to Cancrinia and its relatives was noted by Poljakov 
(1959), who placed several of the present discoid Trichanthe- 
mis species in Cancrinia. He also defined Trichanthemis as 
discoid and removed the radiate species to a new genus 
Glossanthis. His treatment has not been followed by later 
authors, e.g. Tzvelev in Flora URSS (Komarov, 1961). 
Tzvelev noted that the radiate T. aurea and T. radiata are 
related to the discoid T. paradoxos and T. karataviensis, 
respectively. He also indicated the possible relationship 
between the problematic Xylanthemum tianshanicum (Pyre- 
thrum tianshanicum) and Trichanthemis, notably T. butkovii. 
Another intergeneric relationship in need of investigation is 
that of Ugamia and Trichanthemis. Some discoid, small- 
leaved species of Trichanthemis may be more closely related 
to Ugamia than to other Trichanthemis species. It appears 
that Trichanthemis may be paraphyletic. Small segregates, 
possibly Ugamia for example, or even larger ones in Tanace- 
tum may have their sister groups within Trichanthemis. 

The list of species is taken from Flora URSS and Flora 
iranica but with one former synonym re-established as a 
species (T. simulans; Pavlov, 1966). 

T. afghanica Podl. 

T. aulieatensis (B. Fedtsch.) H. Kraschen. 

*T. aurea H. Kraschen. 

*r. butkovii Kovalevsk. 

T. karataviensis Regel & Schmalh. 

*T. litwinowii (H. Kraschen) Tzvelev 

T. paradoxos (Winkler) Tzvelev 

T. radiata H. Kraschen. & Vved. 

* T. simulans (Pavlov) Pavlov 



98 



K. BREMER AND C. J. HUMPHRIES 



10. UGAMIA Pavlov in Vest. Akad. Nauk Kazakh. 
SSR 8:25 (1950). Type species: U. trichanthemoides 
Pavlov {V. angrenica (H. Kraschen.) Tzvelev). 

A compact, tomentose, basally much woody half-shrub. 
Leaves alternate to rosulate, densely set, small, pectinate. 
Capitula solitary on short, nodding peduncles, discoid. 
Involucral bracts with dark brown margins. Receptacle 
almost flat, epaleate. Corolla 5-lobed. Cypselas 10-15- 
ribbed, densely pilose; hairs subulate, with a few basal cells 
and one long apical cell. Pappus of 10-15 white linear scales 
roughly as long as the corolla. 

Distribution. C. Asia. - Monotypic. 

Ugamia angrenica is very characteristic with its compact and 
woody habit, small, pectinate leaves and campanulate 
capitula on short, nodding peduncles. Though discoid as in 
Cancrinia and Cancriniella, it is possibly most closely related 
to part of Trichanthemis , which also has discoid representa- 
tives, with densely pilose cypselas. 

11. RICHTERIA Karelin & Kir. in Bull. Soc. Nat. 
Moscou 15:126 (1842). Type species: R. 
pyrethroides Karelin & Kir. 

Tomentose half-shrubs, basally woody with stems covered in 
sheathing leaf bases. Leaves alternate and basally rosulate, 
pinnatisect. Capitula solitary, pedunculate, radiate. Involu- 
cral bracts with dark brown margins. Receptacle convex, 
epaleate. Ray florets female; Hmb white. Disc corolla 
5-lobed. Cypselas faintly 6-10-ribbed, with sessile glands. 
Pappus of 6-10 obovate, apically brownish scales at least half 
as long as the corolla. 

Distribution. Iran, Afghanistan, C. Asia, Mongolia, China 
in Xinjiang and Tibet, Himalaya. - 3 spp. 

Richteria is commonly treated as part of Pyrethrum (i.e. 
Tanacetum s.l.), but since it was originally described as a 
genus we find it suitable to retain it as such and improve the 
circumscription of Tanacetum (incl. Pyrethrum) by restricting 
it to those species with a short coroniform or auriculate 
pappus. 

Richteria approaches Trichanthemis in habit; the somewhat 
pedunculoid stems are basally covered in more or less sheath- 
ing leaf bases and the leaves are mainly basally arranged. This 
appears to be a plesiomorphic condition within the tribe, 
since a similar habit is shown by representatives of the 
outgroup. The pappus scales are apically brownish as in 
Cancrinia and Allardia, though in the latter genus they are 
more narrow and numerous. Being white-rayed rather than 
discoid or blue-rayed, Richteria is plesiomorphic and appears 
undefined compared to the latter two genera. 

Two species of Richteria have been described and one more 
is added here but there may be more species hidden within 
Tanacetum. 

R. djilgense (Franchet) Bremer & Humphries, comb. nov. 

Basionym: Chrysanthemum djilgense Franchet in Bull. 

Mus. Hist. nat. Paris 2:345 (1896) {Pyrethrum djilgense 

(Franchet) Tzvelev, Tanacetum djilgense (Franchet) 

Podl.). 
R. leontopodium Winkler 
R. pyrethroides Karelin & Kir. (Pyrethrum arassanicum 

(Winkler) O. & B. Fedtsch., Pyrethrum neglectum Tzvelev, 



Pyrethrum pyrethroides (Karelin & Kir.) B. Fedtsch. & H. 
Kraschen., Pyrethrum transiliense (Herder) Regel «fe 
Schmalh., Tanacetum pyrethroides (Karelin & Kir.) 
Schultz-Bip.) - Note: Synonymy after Kovalevskaja (in 
Vvedensky, 1962: 133) and Podlech (in Podlech et al., 
1986). 

12. ALLARDIA Decne in Jacquemont, Voy. Inde 4: 
87 (1842-7). Type species: A. tomentosa Decne. - 
Waldheimia Karelin & Kir. 

Glabrous to densely tomentose perennial herbs. Leaves alter- 
nate to rosulate, densely set, pinnatifid. Capitula solitary, 
pedunculate, radiate. Involucral bracts with dark brown 
margins. Receptacle convex, epaleate. Ray florets female, 
fertile or sterile, or neuter; limb white, pink, or bluish-violet. 
Disc corolla 5-lobed, with a yellow or bluish-violet limb. 
Cypselas faintly 5-10-ribbed, generally with sessile glands, 
sometimes pilose. Pappus of many bristle-like, subulate, 
apically brownish scales as long as or longer than the corolla. 

Distribution. Afghanistan, C. Asia, Mongolia, China in 
Sinkiang and Tibet, and Himalaya. - 8 spp. 

With its white to pink and blue-violet florets and cypselas 
with pappus bristles Allardia may seem out of place in the 
Anthemideae but it is clearly a member of this tribe. The 
pappus 'bristles' are subulate, much elongated scales, similar 
to those of Ugamia (in shape and number) and Richteria and 
Cancrinia (in being apically brownish). Several narrow 
bristles are probably plesiomorphic, and apically brownish 
bristles are probable synapomorphies for Allardia and the 
latter two genera. Tzvelev in Flora URSS (Komarov, 1961) 
also stated that the genus is related to Richteria. 

The publication dates of the various parts of Jacquemont's 
Voyage dans I'lnde are still in dispute, but it appears that pp. 
1-88 of volume 4 were published before 1842 (Stafleu & 
Cowan, 1979). Allardia is then prior to Waldheimia, which 
was published in 1842. Allardia was typified by Tzvelev in 
Flora URSS (Komarov, 1961). He used Waldheimia for the 
section name and Waldheimia tomentosa (Allardia tomen- 
tosa) as the type species. 

*A. huegelii Schultz-Bip. (Waldheimia huegelii (Schultz-Bip.) 
Tzvelev) 

A. lasiocarpa (G. X. Fu) Bremer & Humphries, comb. nov. 
Basionym: Waldheimia lasiocarpa G. X. Fu in Shih & Fu, 
Acta phytotax. sin. 17: 113 (1979). 

A. nivea Hook. f. & Thomson ex C. B. Clarke (Waldheimia 
nivea (C. B. Clarke) Regel) 

*A. stoliczkae C. B. Clarke (Waldheimia stoliczkae (C. B. 
Clarke) Ostenf.) 

A. tomentosa Decne (Waldheimia tomentosa (Decne) Regel, 
Tanacetum tomentosum (Decne) Muradyan) 

A. transalaica (Tzvelev) Bremer & Humphries, comb. nov. 
Basionym: Waldheimia transalaica Tzvelev in Komarov, Fl. 
URSS 26: 875 (1961). 

A. tridactylites (Karelin & Kir.) Schultz-Bip. (A. glabra 
Decne, Tanacetum glabrum (Decne) Muradyan, Waldhe- 
imia glabra (Decne) Regel, Waldheimia tridactylites Kare- 
lin & Kir.), Note: A. glabra was reduced by Kovalevskaja 
(in Vvedensky, 1962:188). 

A. vestita Hook. f. & Thomson ex C. B. Clarke (Waldheimia 
vestita (C. B. Clarke) Pampan.) 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



99 



13. CANCRINIA Karelin & Kir. in Bull. Soc. Nat. 
Moscou 15: 124 (1842). Type species: C. 
chrysocephala Karelin & Kir. 

Compact tomentose perennial herbs. Leaves alternate to 
rosulate, densely set, pinnatifid. Capitula solitary, peduncu- 
late, discoid. Involucral bracts with dark brown margins. 
Receptacle convex, epaleate. Corolla 5-lobed with a narrow 
tube and a distinct limb. Cypselas faintly 7-9-ribbed, glabrous 
or sparsely pilose. Pappus of 5-12 lanceolate, apically brown- 
ish scales as long as or slightly longer than the corolla. 

Distribution. C. Asia, Mongolia and China in Xinjiang. - 4 
spp. 

Cancrinia, originally described as monotypic, was expanded 
to include some 20 species, mainly from Tanacetum, by 
Poljakov (1959) and Tzvelev in Flora URSS (Komarov, 
1961). Tzvelev divided the genus into four sections. Most of 
the species do not belong together with the type species, C. 
chrysocephala and its sister species, C. tianshanica. Tzvelev's 
sect. Polychrysum and sect. Tanacetopsis were elevated to 
genera by Kovalevskaja (in Vvedensky, 1962). The mono- 
typic section Matricarioides (C discoidea) is transferred to 
Matricaria. Cancrinia and Allardia are possibly more closely 
related than indicated by the cladogram. They are similar in 
habit and cypsela morphology, though Allardia has more and 
narrower pappus scales. 

C. chrysocephala Karelin & Kir. 

*C. krasnoboroviiV . Khan. 

C. pamiralaica (Kovalevsk.) Kovalevsk. 

*C. tianshanica (H. Kraschen.) Tzvelev 

14. CANCRINIELLA Tzvelev in Komarov, Fl. URSS 
26:876 (1961). Type species: C. krascheninnikovii 
(Rubtzov) Tzvelev. 

A compact, hirsute, basally woody half-shrub. Leaves alter- 
nate to rosulate, densely set, pectinate. Capitula solitary, 
long-pedunculate, discoid. Involucral bracts in 1-2 rows, 
rather wide, subequal. Receptacle convex, sparsely pilose, 
epaleate. Corolla 5-lobed. Cypselas 10-ribbed. Pappus a 
corona of scales. 

Distribution. C. Asia. - Monotypic. 

This monotypic genus is supposed to be related to Cancrinia. 
Poljakov (1959) stated that it is related to Trichanthemis 
karataviensis, and he placed both species in Cancrinia. 

3. 'TANACETINAE' Bremer & Humphries, 
subtrib. nov. 

Type species: Tanacetum vulgare L. 

Herbae perennes vel frutices vel raro herbae annuae. Cypse- 
lae plerumque oblongae et plus quam quinquecostatae. Pap- 
pus coroniformis e squamis distinctis vel auricula adaxiali 
formatus, vel nuUus. 

Perennial herbs or shrubs, rarely annuals. Leaves alternate or 
sometimes rosulate, generally pinnatisect, rarely entire or 
few-lobed. Capitula solitary or corymbose, generally pedun- 
culate, radiate, disciform or discoid. Involucral bracts some- 
times with dark brown margins. Receptacle flat to conical, 
epaleate or rarely paleate (Heliocauta). Ray floret limb 



white, yellow or pink. Disc corolla 5-lobed. Cypselas often 
oblong and more than 5-ribbed, rarely dorsiventrally flat- 
tened (Heliocauta), sometimes with sessile glands and myxo- 
genic cells. Pappus a corona, or of separate scales, or an 
adaxial auricle, or absent. 

Distribution (Table 9). C. Asia but also in N. America and 
N. Africa, some Tanacetum species introduced in the S. 
hemisphere. - 7 genera, 213 spp. 

The genera of this subtribe, 'Tanacetinae', have no synapo- 
morphies in common. The subtribe is probably paraphyletic 
(see for example, Schultz Bipontinus, 1844fl), and hence put 
within inverted commas following the convention suggested 
by Patterson & Rosen (1977) and Wiley (1981). There is still 
a lot of work to be undertaken on the classification of 
Anthemideae, and Tanacetum particularly is a key genus of 
the tribe. At the present state of knowledge we have felt it 
necessary to adopt a provisional subtribe comprising Tanace- 
tum and a number of odd genera apparently related to this 
genus. 

There are several segregate genera and groups of genera, 
possibly even whole subtribes, which are related to parts of 
Tanacetum, which is thus paraphyletic as presently circum- 
scribed. Apparent examples of such genera are those classi- 
fied in this subtribe. Subtribes which probably have their 
sister groups within Tanacetum include Gonosperminae, 
Handeliinae, and Artemisiinae. The remaining subtribes, 
excluding Ursiniinae and Cancriniinae, may form one group 
(as indicated by the cladogram) or a number of groups also 
with their sister groups within Tanacetum. In fact, Tanacetum 
is a paraphyletic group basal to large parts of the tribe. The 
cladogram shown here is the single most parsimonious one 
derivable from the present data matrix, but the picture may 
be quite different when Tanacetum is resolved into smaller 
monophyletic units. The whole complex is in need of detailed 
investigation, both on the generic and the specific level. 

Clades and characters - Fig. 4, Tables 2, 10. 



f=Tal:j(= 15 Tanacetum 

[= 16 Opisthopappus 

fp= 17 Tanacetopsis 
I— T=.oJ r— 18 Xylanthemum 

L.ra3JL_ 19 Lepidolopha 



=Ta2 



L 



E20 Hippolytia 
21 Heliocauta 



lt=Ta4 

Fig. 4 Cladogram of the 'Tanacetinae' produced by the ie option in 
Hennig86. Cladogram length = 16, consistency index = 100, 
retention index = 100. 

Provisional group Tal - subtribe 'Tanacetinae' 

There is no autapomorphy for this group, but as explained 
above Tanacetum probably contains the sister groups of the 
other genera. 

Tanacetum 

175 Embryo sac tetrasporic. The 10 Tanacetum species inves- 
tigated embryologically (Harling 1951; see discussion under 
Tanacetum) are all tetrasporic. The other genera of 'Tanace- 
tinae' have not been investigated. 

Opisthopappus 

163 Pappus of separate, mainly abaxial, subulate scales. 



100 



K. BREMER AND C. J. HUMPHRIES 



Table 9 General distribution of Tanacetinae and genera. x=indigenous, o=introduced. 





N.Am. 


Eur- 
Asia 


C.&E. 
Asia 


SW 

Asia 


S.Eur. 


N.Afr. 


S.Afr. 


Austr. 
N.Zeal. 


S.Am, 


Tanacetinae 


X 


X 


X 


X 


X 


X 











Tanacetum 


X 


X 


X 


X 


X 


X 


o 


o 


o 


Opisthopappus 

Tanacetopsis 

Xylanthemum 

Lepidolopha 

Hippolytia 

Heliocauta 




X 


X 
X 
X 
X 
X 


X 




X 









Table 10 Data matrix for the Tanacetinae. 1 = presence, = absence, ? = missing data or not applicable, p = polymorphic but scored as the 
plesiomorphic condition, a = polymorphic but scored as the apomorphic condition. 

1111111 1 1111 11 11 1 

1450576 763 633457294013 7515555566176667 21 11236 
5327181 5352678412925048 603356790751568044618919162 

15. Tanacetum alaalll lOpOpOOpOOpOOOOO ??00000000000000ppppppppppp 

16. Opisthopappus alaal?l 7100000000000000 ???000000000000000000000000 

17. Tanacetopsis al?al?l ?010000pOOpOOOOO ????????????0000p0000000000 

18. Xylanthemum alaal?l ?0all00000000000 ???000000000000000000000000 

19. Lepidolopha al?al?l ?011011000p00000 ???????????????00p000000000 

20. Hippolytia al?al?l ?010?00aalal0000 ????????????????00p00000000 

21. Heliocauta al?al?l ?010?00aal00alll ?0??????????????00000000000 



Clade Ta2 

35 Capitula discoid. For reasons of parsimony the five genera 
of this clade are grouped together, though they probably have 
different sister groups within Tanacetum. The radiate species 
Xylanthemum tianshanicum is probably misplaced in this 
genus. 

Tanacetopsis 

There is no autapomorphy for this genus. 

Clade Ta3 

2 Plants shrubby. 

Xylanthemum 

166 Pappus adaxially long. 

Lepidolopha 

37 Involucre rather narrowly urceolate. 

38 Involucral bracts in 4-7 rows. 

Clade Ta4 

44 Involucral bracts with dark brown margins. This character 
is present also in several Tanacetum and some Tanacetopsis 
species. 

51 Floral parts with resin canals. Floral resin canals also occur 
in scattered species of the other genera of the subtribe. 

172 Pappus absent in ray and disc cypselas. 

Hippolytia 

29 Capitula densely corymbose. Corymbose capitula are com- 
mon in Tanacetum and present also in some species of 
Tanacetopsis and Lepidolopha. Some specialized alpine spe- 
cies of Hippolytia have solitary capitula. 

92 Anthers caudate. 



Heliocauta 

45 Receptacle paleate. 

100 Style immersed in a lobed nectary. 

114 Cypselas dorsiventrally flattened. 

138 Cypselas with scattered elongated resin sacs. 

15. TANACETUM L., Sp. pi: 1028 (1753). Type 
species: T. vulgare L. - Balsamita Miller - 
Gymnocline Cass. - Hemipappus K. Koch - 
Pyrethrum Zinn - Spathipappus Tzvelev. 

Perennial, rarely annual herbs or seldom small, basally 
woody half-shrubs. Leaves alternate or rarely rosulate, pin- 
natifid to pinnatisect or rarely entire, serrate. Capitula soli- 
tary or corymbose, radiate, disciform, or discoid. Involucral 
bracts sometimes with a dark brown margin. Receptacle flat 
or convex, epaleate. Ray florets female or neuter; limb 
yellow, white, or pink. Disc corolla 5-lobed. Cypselas gener- 
ally oblong, 5-12-ribbed, often with sessile glands, never with 
myxogenic cells. Pappus a short or well developed corona, 
sometimes of short free scales, rarely an adaxial auricle (very 
rarely absent). Embryo sac tetrasporic. 

Distribution. Europe and temperate Asia, also in N. 
Africa {T. annuum, T. corymbosum) and several species in 
N. America, some species introduced in the S. hemisphere. - 
152 spp. 

Tanacetum s. 1. is a large and poorly understood boreal genus 
with many little-known Asian representatives. Variation in 
habit, foHage, inflorescences, and capitula structure is consid- 
erable. As presently circumscribed, the genus may be recog- 
nized by the epaleate receptacle and the non-myxogenic, 
generally straight and ribbed cypselas with a coroniform or 
rarely auriculate pappus. Occasional species with epappose 
cypselas currently placed in Tanacetum are probably mis- 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



101 



placed and are likely to be transferred to other genera. 

One example of a probable misplaced species is T. micro- 
phyllum, with triangular apical anther appendages and thin- 
walled, obovoid cypselas with minute pappus scales and no 
ribs. These characters are reminiscent of subtribe Artemisii- 
nae, where this species probably should be accommodated. 

Pyrethrum differs from Tanacetum s. s. by its white or pink 
rays. Tanacetum s. s. has discoid or disciform capitula or, if 
radiate, with yellow rays. Tzvelev (in Komarov, 1961) diag- 
nosed Pyrethrum as white-rayed and Tanacetum as yellow- 
rayed or disciform (heterogamous). He classified discoid 
(homogamous) species in Cancrinia. Tzvelev also noted the 
heterogeneous nature of Pyrethrum and speculated that Can- 
crinia (s. 1.) evolved from Pyrethrum by loss of rays, where- 
after Tanacetum (s. s.) evolved from Cancrinia by 
transformation of the outer, yellow disc florets into short, 
yellow rays. Except for the often short lamina the yellow rays 
of Tanacetum (s. s.) are similar to the white or pink ones of 
Pyrethrum. In other Anthemideae, e.g. Cotula, where pseu- 
dorays have evolved from the disc florets, their discoid nature 
is clearly recognized. Also ray colour is very homoplasious 
within the tribe, many well-defined genera having species 
with both white and yellow rays. The relationship between 
Pyrethrum and Tanacetum s. s. is close and involves several 
sister group relationships between species and groups of 
species currently classified in the two genera. We follow 
Heywood in Flora europaea (Tutin et al., 1976; also Hey- 
wood, 1954) and Grierson in Flora of Turkey (Davis, 1975) in 
uniting the two genera. 

Tzvelev (in Komarov, 1961) recognized 14 sections of 
Pyrethrum and four sections of Tanacetum. Sections Leucan- 
themopsis and Pyrethrum section Richteria are considered by 
us as separate genera in different subtribes. Section Balsamita 
(T. balsamita) is sometimes considered as a separate genus 
distinguished by its entire, serrate leaves. Harling (1951) in 
describing the unusual tetrasporic embryo sac development 
and Favarger (1966) referring to chemistry both recom- 
mended that Balsamita should be kept separate from Tanace- 
tum. The actual presence of a tetrasporic embryo sac in 
Balsamita seems to strengthen its position within Tanacetum 
(together with the type species T. vulgare and others), though 
there is a difference in embryo sac development. Chemical 
evidence is far too scattered to support a removal of Bal- 
samita from Tanacetum. In morphology Balsamita is similar 
to several representatives of Pyrethrum sensu Tzvelev and we 
follow him and Grierson in Flora of Turkey (Davis, 1975) in 
reducing Balsamita to synonymy. 

Tzvelev's other sections of Pyrethrum and Tanacetum are a 
mixture of isolated and, in most cases, vaguely defined 
entities. Pyrethrum sect. Trichanthemopsis with the single 
species P. tianshanicum has been transferred to Xylanthe- 
mum. Other more or less isolated sections are the woody 
Pyrethrum sect. Xylopyrethrum and Tanacetum sect. Asterot- 
richa. The woody sections, subtribe Cancriniinae, and the 
shrubby genera Xylanthemum and possibly also Lepidolopha 
may be more closely related than indicated by the present 
classification. 

Harling (1951) reported tetrasporic embryo sacs in all 10 
species of Tanacetum so far investigated {T. balsamita, T. 
camphoratum, T. cinerariifolium, T coccineum (Chrysanthe- 
mum marschallii) , Tanacetum corymbosum, T macrophyl- 
lum, T. millefolium, T. parthenium, T. roseum and T. 
vulgare. Interestingly, these species represent nine of 



Tzvelev's different sections, but not the more isolated ones 
mentioned above. 

Tzvelev accepted only heterogamous (disciform or yellow- 
rayed) species in Tanacetum s. s. Thus the discoid T. argen- 
teum was accommodated in the genus Hemipappus, further 
distinguished by its auriculate pappus. On other characters 
Hemipappus fits well into Tanacetum and it was not accepted 
as a genus by Grierson in Flora of Turkey, who included not 
only T. argenteum but also some related Turkish endemic 
species (T. depauperatum, T. haradjanii, T. tomentellum) in 
Tanacetum. This solution is followed here, and we include 
other species with an auriculate pappus in Tanacetum (for- 
merly Spathipappus, discussed below). Except for the pappus 
there are no indications that Hemipappus and Spathipappus 
are closely related sister groups. 

Spathipappus described by Tzvelev in Flora URSS, is 
supposed to differ from Pyrethrum by its auriculate pappus 
and sterile rays. Muradyan (1970) found that the fruits are 
very similar to those of other Tanacetum species and trans- 
ferred S. griff ithii to Tanacetum. The other two species, S. 
chitralensis and S. porphyrostephanus , are recombined here. 

The list of species is compiled mainly from Flora europaea 
(Tutin et al., 1976), Flora of Turkey (Davis, 1975), and Flora 
URSS (Komarov, 1961, including Pyrethrum) with species 
described later or from other areas added. 

T. abrotanifolium (L.) Druce. Turkey, Caucasus, Iran. 

T. abrotanoides Bremer & Humphries, nom. nov. Basionym: 
Pyrethrum abrotanifolium Bunge ex Ledeb., Fl. ross. 2: 
549 (1845) - Note: In Tanacetum the epithet abrotanifolium 
is already occupied by the preceding species in the list. C. 
Asia, China in Xinjiang. 

T. achilleifolium (M. Bieb.) Schultz-Bip. E. Europe, C. Asia. 

T. akinfievii (Alexej.) Tzvelev. Caucasus. 

T. alatavicum Herder {Pyrethrum alatavicum (Herder) O. & 
B. Fedtsch.). C. Asia, Mongolia, China in Xinjiang. 

T. albipannosum Huber-Mor. & Grierson. Turkey. 

T. alyssifolium (Bornm.) Grierson. Turkey. 

T. annuum L. SW Europe in France, Spain, and Portugal, N. 
Africa in Morocco. 

*T. arctodzhungaricum (Golosk.) Bremer & Humphries, 
comb. nov. Basionym: Pyrethrum arctodzhungaricum 
Golosk. in Bot. Mater. Gerb. Inst. Bot. Akad. Nauk 
Kazakh. SSR 7: 35 (1971). C. Asia. 

T. archibaldii Podl. Iran. 

T. argenteum (Lam.) Willd. {Hemipappus argenteus (Lam.) 
Tzvelev, Hemipappus canus K. Koch). Turkey, Middle 
East, Caucasus. 

*T. argyranthemoides (Boiss. & Kotschy) Schultz-Bip. Iran. 

T. armenum (DC.) Schultz-Bip. {Pyrethrum heldreichianum 
Fenzl ex Tchich.). Turkey. 

T. aromaticum (Tzvelev) Bremer & Humphries, comb. nov. - 
Basionym: Pyrethrum aromaticum Tzvelev in Komarov, Fl. 
URSS 26: 222 (1961). - Note: the name Pyrethrum aromati- 
cum must be considered a new species described by 
Tzvelev, and not a new combination based on Tripleuro- 
spermum aromaticum Rupr. ex Boiss. as stated by Tzvelev. 
That name was only published pro syn. by Boissier (1875: 
334). Caucasus. 

*T. artemisioides Schultz-Bip. in Hook. f. Himalaya. 

*T. atkinsonii (C. B. Clarke) Kitam. {Pyrethrum atkinsonii 
(C. B. Clarke) Ling & Shih). China, Himalaya. 

T. aucheranum (DC.) Schultz-Bip. (Pyrer/zrum aucheranum 
DC). Turkey, Caucasus. 



102 



K. BREMER AND C. J. HUMPHRIES 



T. aucheri DC. Turkey, Syria, Lebanon, Israel. 

T. audibertii (Req.) DC. Sardinia. 

T. balsamita L. {Balsamita major Desf., Pyrethrum balsamita 
(L.) Willd., Pyrethrum balsamitoides (Nab.) Tzvelev, Pyre- 
thrum majus (Desf.) Tzvelev). SW Asia from Turkey, 
Caucasus, Iran, and Afghanistan, also widely cultivated 
and naturalized. 

T. bamianicum Podl. Afghanistan. 

*T. bardayanum DC. {T. turlanicum (Pavlov) Tzvelev). C. 
Asia, China in Xinjiang. 

T. bipinnatum (L.) Schultz-Bip. NE Europe, Siberia, Alaska. 

T. boreale Fischer ex DC. Siberia, C. Asia and Far East, 
China, Korea, Alaska, Canada. 

T. budjurnense (Rech. f.) Tzvelev. Iran. 

*T. cadmeum (Boiss.) Heyw. Turkey. 

T. camphoratum Less. Western N. America from British 
Columbia S. to California. 

T. canescens DC. {T. gilliatii (Turrill) Parsa, T. modestum 
(Heimerl ex Stapf) Parsa). Turkey, Caucasus, Iran. 

T. cappadocicum (DC.) Schultz-Bip. Turkey. 

T. changaicum (H. Kraschen. ex Grubov) Bremer & 
Humphries, comb. nov. Basionym: Pyrethrum changaicum 
H. Kraschen. ex Grubov in Bot. Mater. Gerb. bot. Inst. V. 
A. Komorova 11: 23 (1955). Mongolia. 

T. chiliophyllum (Fischer & C. Meyer) Schultz-Bip. {T. 
heimerli (Nab.) Parsa, T. oUgocephalum (DC.) Schultz- 
Bip.). Turkey, Caucasus, Iraq, Iran. 

T. chitralense (Podl.) Bremer & Humphries, comb. nov. 
Basionym: Spathipappus chitralensis Podl., Fl. iranica 158: 
151 (1986). Pakistan. 

T. ciUcium (Boiss.) Grierson. Turkey, Middle East, Iraq. 

T. cinerariifoUum (Trevir.) Schultz-Bip. {Pyrethrum cinerarii- 
folium Trevir). SE and E. Europe, Caucasus, C. Asia, 
China, often cultivated as the source of the insecticide 
pyrethrin. 

T. coccineum (Willd.) Grierson {Pyrethrum coccineum 
(Willd.) Vorosch., Pyrethrum chamaemelifolium (Sommier 
& Levier) Sosn.). Turkey, Caucasus, Iran, often cultivated 
as the garden 'Pyrethrum' . 

T. corymbiforme (Tzvelev) Bremer & Humphries, comb, 
nov. Basionym: Pyrethrum corymbiforme Tzvelev in 
Komarov, Fl. URSS 26: 873 (1961). C. Asia, China in 
Xinjiang. 

T. corymbosum (L.) Schultz-Bip. {Pyrethrum corymbosum 
(L.) Willd., Pyrethrum clusii Fischer ex Reichb.). N. 
Africa in Morocco and Algeria, most of Europe except N. 
parts, Turkey, Caucasus, C. Asia. 

*T. crassipes (Stchegl.) Tzvelev. C. Asia, China in Xinjiang. 

T. daghestanicum (Rupr. ex Boiss.) Bremer & Humphries, 
comb. nov. Basionym: Chamaemelum daghestanicum 
Rupr. ex Boiss., Fl. orient. 3: 334 (1875) {Pyrethrum 
daghestanicum (Rupr. ex Boiss.) Rupr. ex Flerov). Cauca- 
sus. 

*T. demetrii (Manden.) Bremer & Humphries, comb. nov. 
Basionym: Pyrethrum demetrii Manden. in Zametki Sist. 
Geogr. Rast. 22: 60 (1961). Caucasus. 

T. densum (Labill.) Schultz-Bip. Turkey. 

T. depauperatum (Post) Grierson. Turkey. 

T. dolomiticum (Galushko) Bremer & Humphries, comb, 
nov. Basionym: Pyrethrum dolomiticum Galushko in 
Novit. Syst. PI. Vase. Acad. Sci. URSS 6: 218 (1970). 
Caucasus. 

*T. dumosum Boiss. Iran. 

T. eginense (Hausskn. ex Bornm.) Grierson. Turkey. 



*T. elongatum (Bornm. & Gauba) Parsa. Iran. 

*T. falcatolobatum H. Kraschen. {Cancrinia maximoviczii 

Winkler). China. 
T. falconeri Hook. f. Himalaya. 

T. ferulaceum (Webb ex Berth.) Schultz-Bip. Canary Islands. 
*T. funkii Schultz-Bip. ex Willk. SW Europe in Spain. 
*T. galae (Popov) Nevski {Pyrethrum galae Popov). C. Asia. 
T. galushkoi (Prima) Bremer & Humphries, comb. nov. 

Basionym: Pyrethrum galushkoi Prima in Nov. Sist. 

Vysshikh Rast. 11: 277 (1974). Caucasus. 
T. germanicopolitanum (Bornm. & Heimerl) Grierson. Tur- 
key. 
T. ghoratense Podl. Afghanistan. 
T. glanduliferum (Sommier & Levier) Bremer & Humphries, 

comb. nov. Basionym: Pyrethrum glanduliferum Sommier 

& Levier, Decas Comp. Nov. Cauc. 87 (1895). Caucasus. 
*T. griffithii (C. B. Clarke) Muradyan {Spathipappus griffithii 

(C. B. Clarke) Tzvelev). Afghanistan, C. Asia, Himalaya. 
T. grossheimii (Sosn.) Muradyan {Pyrethrum grossheimii 

Sosn.). Caucasus, Iran. 
T. haradjanii (Rech. f.) Grierson. Turkey. 
*T. haussknechtii (Bornm.) Grierson. Turkey. 
*T. herderi Regel & Schmal. SW Asia. 
*T. heterotomum (Bornm.) Grierson. Turkey. 
T. hissaricum (H. Kraschen.) Bremer & Humphries, comb. 

nov. Basionym: Pyrethrum hissaricum H. Kraschen. in 

Feddes Reprium 26: 26 (1929). C. Asia. 
T. hololeucum (Bornm.) Podl. Iran. 
T. huronense Nutt. N. America in Alaska, Canada and 

Michigan. 
T. karelinianum Bremer & Humphries, nom. nov. Basionym: 

Pyrethrum karelinii H. Kraschen. in Nov. Sist. Vysshikh 

Rast. 9: 157 (1946). C. Asia. 
*T. karelinii Tz\e\e\ . C. Asia. 
T. kaschgarianum Bremer & Humphries, nom. nov. 

Basionym: Pyrethrum kaschgaricum H. Kraschen. in Nov. 

Sist. Vysshikh Rast. 9: 158 (1946) (non T. kaschgaricum H. 

Kraschen.). China. 
T. kelleri (Krylov & Plotn.) Takht. {Chrysanthemum kelleri 

Krylov & Plotn., Pyrethrum kelleri (Krylov & Plotn.) H. 

Kraschen.). C. Asia. 
T. khorassanicum (H. Kraschen.) Parsa {T. czerniakowskae 

(H. Kraschen.) Parsa). Iran. 
T. kotschyi (Boiss.) Grierson {Pyrethrum kotschyi Boiss.). 

Turkey, Caucasus, Iraq, Iran. 
T. krylovianum (H. Kraschen.) Bremer & Humphries, comb. 

nov. Basionym: Pyrethrum krylovianum H. Kraschen. in 

Bot. Mater. Gerb. bot. Inst. V. A. Komorova 9: 155 (1946). 

C. Asia, China in Xinjiang. 
*T. kubense (Grossh.) Muradyan {Pyrethrum kubense 

Grossh.). Caucasus. 
*T. lanuginosum Schultz-Bip. & Herder {Pyrethrum lanugi- 

nosum (Schultz-Bip. & Herder) Tzvelev). E. Siberia, Mon- 
golia. 
T. leptophyllum (Steven) Schultz-Bip. {Pyrethrum leptophyl- 

lum Steven). Caucasus. 
*T. longipedunculatum (Sosn.) Tzvelev. Caucasus. 
*T. macrocephalum Pampan. Himalaya. 
T. macrophyllum (Waldst. & Kit.) Schultz-Bip. {Pyrethrum 

macrophyllum (Waldst. & Kit.) Willd.). C. and SE 

Europe, Turkey, Caucasus. 
T. marionii (Albov) Bremer & Humphries, comb. nov. 

Basionym: Pyrethrum marionii Albov in Bull. Herb. Bois- 
siere. 92 (1895). Caucasus. 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



103 



T. maymanense Podl. Afganistan. 

T. microphyllum DC. SW Europe in Spain and Portugal. 

*T. mikeschinii (Tzvelev) Takht. (Pyrethrum mikeschinii 
Tzvelev). C. Asia. 

T. millefolium (L.) Tzvelev (7. kittaryanum (C. Meyer) 
Tzvelev). E. Europe, S. Siberia, C. Asia. 

*T. mucroniferum Huber-Mor. & Grierson. Turkey. 

T. mucronulatum (Hoffsgg & Link) Heyw. SW Europe in 
Portugal. 

T. nitens (Boiss. & Noe) Grierson. Turkey. 

T. nivale Schultz-Bip. Iraq. 

T. niveum (Lagasca) Schultz-Bip. {Pyrethrum fruticulosum 
Biehler). Caucasus. 

T. nuristanicum Podl. Afghanistan. 

*T. odessanum (Klokov) Tzvelev. E. Europe. 

T. oltense (Sosn.) Grierson. Turkey. 

T. ordubadense (Manden.) Bremer & Humphries, comb, 
nov. Basionym: Pyrethrum ordubadense Manden. in A^ov. 
Sist. Vysshikh Rast. 19: 358 (1959). Caucasus. 

T. oxylepis (Bordz.) Grierson. Caucasus, Turkey. 

T. oxystegium (Sosn.) Grierson. Turkey. 

*T. paczoskii (Zef.) Tzvelev. Krym. 

T. pakistanicum Podl. Pakistan. 

T. paleaceum Podl. Afghanistan. 

*7'. paradoxum Bornm. Iran. 

T. parthenifolium (Willd.) Schultz-Bip. (Pyrethrum partheni- 
folium Willd.). SW Asia from Turkey to Caucasus, Iran, 
Aghanistan, and C Asia. 

T. parthenium (L.) Schultz-Bip. {Pyrethrum parthenium (L.) 
Smith). N. Africa, SE and E. Europe, SW Asia from 
Turkey to Caucasus, Iran, Aghanistan, and C. Asia, also 
widely naturalized. 

*T. petiolosum Pampan. Himalaya. 

T. petrareum (Shih) Bremer & Humphries, comb. nov. 
Basionym: Pyrethrum petrareum Shih in Bull. hot. Lab. 
n.-eastFor. Inst. 6: 10 (1980). China. 

*T. peucedanifolium (Sosn.) Bremer & Humphries, comb, 
nov. Basionym: Pyrethrum parthenifolium Willd. var. peu- 
cedanifolium Sosn. in Trudy tiflis. bot. Sada 17: 35 (1915) 
{Pyrethrum peucedanifolium (Sosn.) Manden.). Caucasus. 

*T. pinnatum Boiss. {T. flavovirens (Boiss.) Tzvelev, T. 
tamrutense (Sosn.) Sosn.). Turkey, Caucasus, Iraq, Iran. 

T. podlechii Bremer & Humphries, nom. nov. Basionym: 
Pyrethrum komarovii Sosn. in Dokl. Akad. Nauk armyan. 
SSR 2 (4): 119 (1945) (non Tanacetum komarovii (Winkler) 
Muradyan). Caucasus. 

T. polycephalum Schultz-Bip. {T. argyrophyllum (K. Koch) 
Tzvelev, T. duderanum (Boiss.) Tzvelev, T. heterophyllum 
Boiss., T. junesarense (Bornm.) Parsa, T. myriophyllum 
Willd.). Turkey, Caucasus, Iraq, Iran. 

T. porphyrostephanum (Rech. f.) Bremer & Humphries, 
comb. nov. Basionym: Chrysanthemum porphyrostepha- 
num Rech. f., Symb. Afghan. 2: 47 (1955) {Spathipappus 
porphyrostephanus (Rech. f.) Tzvelev). Iran. 

T. poteriifolium (Ledeb.) Grierson {Pyrethrum poteriifolium 
Ledeb.). Turkey, Caucasus. 

T. praeteritum (Horw.) Heyw. Turkey. 

T. pseudoachillea Winkler. C. Asia. 

T. ptarmiciflorum (Webb & Berth.) SchuUz-Bip. Canary 
Islands. 

*T. pulchellum (Turcz.) Schultz-Bip. {Pyrethrum pulchellum 
Turcz.). E. Siberia. 

T. pulchrum (Ledeb.) Schultz-Bip. {Pyrethrum pulchrum 
Ledeb.). C. Asia, Mongolia, China in Xinjiang. 



T. punctatum (Desr.) Grierson {Pyrethrum punctatum 
(Desr.) Bordz. ex Sosn.). Turkey, Caucasus. 

*T. richterioides (Winkler) Bremer & Humphries, comb, 
nov. Basionym: Chrysanthemum richterioides Winkler in 
Trudy imp. S.-Peterb. bot. Sada 10: 86 (1887) {Pyrethrum 
richterioides (Winkler) H. Kraschen.). China. 

*T. robustum Hook. f. & Thomson ex C. B. Clarke. Hima- 
laya. 

T. roseum (Adams) Schultz-Bip. {Pyrethrum roseum 
(Adams) M. Bieb.). Caucasus. 

T. roylei (DC.) Podl. W. Himalayas. 

T. salsugineum Podl. Iran. 

*T. sanguineum (Parsa) Parsa. Iran. 

T. santolina Winkler. S. European USSR, S. Siberia. C. 
Asia, China in Xinjiang. 

*T. saxicolum (H. Kraschen.) Tzvelev. C. Asia. 

T. sclerophyllum (H. Kraschen.) Tzvelev. S. European 
USSR. 

*T. scopulorum (H. Kraschen.) Tzvelev. C. Asia, China in 
Xinjiang. 

T. semenovii Herder {Pyrethrum semenovii (Herder) Winkler 
ex O. & B. Fedtsch.). C. Asia. 

T. sericeum (Adams) Schultz-Bip. {Pyrethrum sericeum 
(Adams) M. Bieb.). Turkey, Caucasus. 

T. sevanense (Sosn.) Bremer & Humphries, comb. nov. 
Basionym: Pyrethrum sevanense Sosn. ex Gross., Fl. Kauk. 
4: 137 (1934). Caucasus, Iran. 

T. silaifolium (Steven) Schultz-Bip. {Pyrethrum silaifolium 
Steven). Caucasus. 

T. silvicola Podl. Afghanistan. 

T. sinaicum (Fresen.) Del. ex Bremer & Humphries, comb, 
nov. Basionym: Santolina sinaica Fresen., Mus. senckenb. 
1: 83 (1833) {Pyrethrum santolinoides DC.) - Note: The 
combination Tanacetum sinaicum has hitherto not been 
validly published; it was only published pro syn. by de 
CandoUe (1837: 59). Middle East in Sinai. 

*T. sipikorense (Bornm.) Grierson {Pyrethrum oxylepis 
(Bordz.) Tzvelev). Turkey, Caucasus. 

T. songaricum (Tzvelev) Bremer & Humphries, comb. nov. 
Basionym: Pyrethrum songaricum Tzvelev in Komarov, Fl. 
URSS 26: 874 (1961). C. Asia. 

T. sorbifolium (Boiss.) Grierson {Pyrethrum sorbifolium 
Boiss.). Turkey, Caucasus. 

*T. stapfianum (Rech. f.) Podl. Iran. 

T. tabrisianum (Boiss.) Sosn. & Takht. Turkey, Caucasus, 
Iran. 

T. tanacetoides (DC.) Tzvelev. C. Asia, Mongolia, China in 
Xinjiang. 

T. tatsienense (Bureau & Franchet) Bremer & Humphries, 
comb. nov. Basionym: Chrysanthemum tatsienense Bureau 
& Franchet in /. Bot. 5: 72 (1891) {Pyrethrum tatsienense 
(Bureau & Franchet) Ling ex Shih). China in Tibet. 

T. tenuisectum (Boiss.) Podl. Iran. 

T. tenuissimum (Trautv.) Gross. Caucasus, Iran, Afghani- 
stan. 

T. tirinense Podl. Afghanistan. 

T. tomentellum (Boiss.) Grierson. Turkey. 

T. trichophyllum (Sosn.) Bremer & Humphries, comb. nov. - 
Basionym: Pyrethrum trichophyllum Sosn. in Zametki Sist. 
Geogr. Rast. 15: 2 (1949) (non Griseb.) {Pyrethrum tri- 
cholobum Sosn. ex Manden.). Caucasus. 

T. trifoliolatum Podl. Iran. 

*T. turcomanicum (H. Kraschen.) Tzvelev. Iran, C. Asia. 

*T. ulutavicum Tzvelev. C. Asia. 



104 



K. BREMER AND C. J. HUMPHRIES 



T. uniflorum (Fischer & C. Meyer ex DC.) Schultz-Bip. 

Turkey, Caucasus, Iran. 
*T. uralense (H. Kraschen.) Tzvelev. E. Europe, S. Siberia, 

C. Asia. 
*T. vahlii DC. SW Europe in Spain. 
T. vulgare L. Throughout most of Europe and temperate 

Asia, introduced in America, AustraHa and New Zealand. 
T. walteri (Winkler) Tzvelev. Iran, C. Asia. 
*T. willkommii Schultz-Bip. SW Europe in Spain. 
*T. yabrudae (Mout.) Charpin & Dittrich {Pyrethrum 

yabrudae Mout.). Middle East in Syria. 
T. zahlbruckneri (Nab.) Grierson. Turkey. 

16. OPISTHOPAPPUS Shih in Acta phytotax. sin. 17: 
110 (1979). Type species: O. taihangensis (Ling) 
Shih. 

Perennial herbs with basally somewhat woody stems. Leaves 
alternate, pinnatisect. Capitula solitary, radiate. Receptacle 
convex to conical, epaleate. Ray florets female, fertile; limb 
white or pinkish, many-veined. Disc corolla 5-lobed, with 
sessile glands. Cypselas obovoid, 3-5-ribbed, thin-walled, 
with myxogenic cells. Pappus of separate, mainly abaxial, 
subulate scales. 

Distribution. NE central China. - 2 spp. 

The position of this genus is unclear. It appears to be a 
derivative of Tanacetum, including Pyrethrum, where the 
type species was formerly accommodated. According to Shih 
it differs from Pyrethrum and Spathipappus (here included in 
Tanacetum) by its thickly ribbed cypselas and by its mainly 
abaxial pappus of separate scales, rather than a corona or an 
auricle. The cypsela ribs as such are not thick but since they 
are invested with large myxogenic cells, dry cypselas appear 
thickly ribbed. The presence of myxogenic cells is indeed a 
character distinguishing Opisthopappus from Tanacetum, 
which has non-myxogenic cypselas. 

O. longilobus Shih 

O. taihangensis (Ling) Shih 

17. TANACETOPSIS (Tzvelev) Kovalevsk. in 
Vvedensky, Fl. uzbekistana 6: 138 (1962). Type 
species: T. mucronata (Regel & Schmalh.) 
Kovalevsk. - Cancrinia sect. Tanacetopsis Tzvelev. 

Perennial herbs or basally woody half-shrubs. Leaves alter- 
nate, pinnatisect. Capitula solitary or generally laxly to 
densely corymbose, discoid. Involucral bracts sometimes with 
dark brown margins. Receptacle convex, epaleate, rarely 
sparsely pilose. Corolla 5-lobed. Cypselas 5-ribbed, some- 
times with sessile glands, generally with myxogenic cells. 
Pappus a corona of free or united scales. 

Distribution. Iran, Afghanistan and C. Asia. -21 spp. 

Tanacetopsis is based on a section of Cancrinia described by 
Tzvelev (1971) and in Komarov (1961). Tzvelev transferred a 
number of Tanacetum and Lepidolopsis species to Cancrinia, 
although he noted that his Cancrinia sections could be treated 
as separate genera. He argued that their relationship was 
shown by similar floret, cypsela, and pollen morphology. The 
type species of Cancrinia (see below) is very different from 
Tanacetopsis species, in habit as well as floret and cypsela 
morphology. The removal of Tanacetopsis from Cancrinia is 



well justified (Kovalevskaja, 1972), but we doubt its status as 
a genus separate from Tanacetum, at least as presently 
circumscribed. 

Tzvelev noted that some species of Cancrinia, i.e. Tanac- 
etopsis, have a sparsely pilose receptacle, a character which 
does not occur in Tanacetum. Since only some species are 
involved, however, it cannot be used as a synapomorphy for 
the genus. The alleged difference in shape of involucre, also 
noted by Tzvelev, campanulate in Cancrinia and Tanacetopsis 
and cyathiform in Tanacetum, breaks down after examination 
of the numerous species involved. 

Tzvelev also noted that his monotypic section Leptanthe- 
mum of Pyrethrum {Tanacetum leptophyllum) is almost iden- 
tical to some species of Cancrinia sect. Tanacetopsis (e.g. the 
type species) except in being radiate rather than discoid. In 
fact, Tanacetopsis differs from Tanacetum simply by homoga- 
mous (discoid) as opposed to heterogamous capitula. With 
the inclusion of Hemipappus in Tanacetum there are also 
discoid representatives of Tanacetum. Podlech (in Podlech et 
al., 1986) included Tanacetopsis as a section of Tanacetum. 
Since we have seen only limited material of Tanacetopsis and 
since this whole Tanacetum group is in need of generic 
revision, we retain Tanacetopsis provisionally as a genus. 

T. afghanica (Gilli) Bremer & Humphries, comb. nov. 

Basionym: Chrysanthemum afghanicum Gilli in Feddes 

Reprium Spec. nov. veg. 68: 93 (1963) {Tanacetum afghani- 
cum (Gilli) Podl.) 
*T. botschantzevii (Kovalevsk.) Kovalevsk. {Cancrinia 

botschantzevii (Kovalevsk.) Tzvelev) 
T. doabensis (Podl.) Bremer & Humphries, comb. nov. 

Basionym: Tanacetum doabense Podl., Fl. iranica 158: 131 

(1986). 
*T. eriobasis (Rech. f.) Kovalevsk. {Tanacetum eriobasis 

(Rech. f.) Kovalevsk.) 
*T. ferganensis (Kovalevsk.) Kovalevsk. {Cancrinia ferganen- 

sis (Kovalevsk.) Tzvelev) 
T. freitagii (Podl.) Bremer & Humphries, comb. nov. 

Basionym: Tanacetum freitagii Podl., Fl. iranica 158: 132 

(1986). 
T. golovskovii (Polj.) Karmysch. {Cancrinia golovskovii 

(Polj.) Tzvelev) 
T. hedgei (Podl.) Bremer & Humphries, comb. nov. 

Basionym: Tanacetum hedgei Podl., Fl. iranica 158: 130 

(1986). 
*T. kamelinii Kovalevsk. 
T. karataviensis (Kovalevsk.) Kovalevsk. {Cancrinia karatav- 

ica Tzvelev) 
* T. kjurendaghii Kurbanov 
*T. krascheninnikovii (Nevski) Kovalevsk. {Cancrinia nevskii 

Tzvelev) 
T. mucronata (Regel & Schmalh.) Kovalevsk. 
*T. pjataeviae (Kovalevsk.) Karmysch. {Cancrinia pjataeviae 

(Kovalevsk.) Tzvelev) 
*T. platyrachis (Boiss.) Kovalevsk. 
T. santoana (H. Kraschen., Popov & Vved.) Kovalevsk. 

{Cancrinia santoana (H. Kraschen., Popov & Vved.) 

Tzvelev) 
*T. setacea (Regel & Schmalh.) Kovalevsk. {Cancrinia seta- 

cea (Regel & Schmalh.) Tzvelev) 
T. submarginata (Kovalevsk.) Kovalevsk. {Cancrinia sub- 

marginata (Kovalevsk.) Tzvelev) 
*T. subsimilis (Rech. f.) Kovalevsk. {Cancrinia subsimilis 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



105 



(Rech. f.) Tzvelev, Tanacetum subsimile (Rech. f.) Kova- 

levsk.) 
*T. tripinnatifida (Oliver) Kovalevsk. 
T. urgutensis (Popov) Kovalevsk. {Cancrinia urgutensis 

(Popov) Tzvelev) 

18. XYLANTHEMUM Tzvelev in Komarov, Fl. URSS 
26: 877 (1961). Type species: X. fisherae (Aitch. & 
Hemsley) Tzvelev. 

Shrublets generally with virgate eventually leaf-less stems, 
sometimes much woody basally. Leaves alternate, pinnati- 
sect. Capitula solitary, pedunculate, discoid or possibly also 
radiate with white rays. Involucral bracts in several rows, the 
outer much smaller than the inner. Receptacle flat, epaleate. 
Corolla 5-lobed. Cypselas oblong, 5-6-ribbed, generally with 
sessile glands and myxogenic cells. Pappus an adaxial auricle 
or a corona of several adaxially more developed scales. 

Distribution. Iran, Afghanistan and C. Asia. - 9 spp. 

According to Tzvelev (in Komorov, 1961) Xylanthemum 
differs from Tanacetum by its homogamous (discoid) 
capitula, flat receptacle, and auriculate pappus. However, 
these characters occur also in Tanacetum with Hemipappus 
and Spathipappus included (see discussion under Tanace- 
tum). Furthermore, X. pamiricum has a pappus of free scales 
developed more strongly on the adaxial side, probably a 
plesiomorphic condition when compared to the auriculate 
pappus in X. fisherae. Thus, the genus as a whole has not a 
pappus of a single auricle but at most, as expressed by 
Tzvelev, an auricle sometimes cleft to the base. 

Nevertheless, it is possible that Xylanthemum as circum- 
scribed by Tzvelev is a monophyletic group defined not only 
by these characters as synapomorphies, but also by its 
shrubby habit and involucral bracts in several rows. Tzvelev 
also stated that the species are closely related geographical- 
ecological races of one natural unit. Muradyan (1970) investi- 
gated fruits of Tanacetum and Xylanthemum and concluded 
that they are different and that Spathipappus belongs in 
Tanacetum rather than together with Xylanthemum, despite 
the similarity in pappus structure. The relationship of Xylan- 
themum to Tanacetum or probably rather to a part of 
Tanacetum, which then becomes paraphyletic by the exclu- 
sion of Xylanthemum, deserves further investigation. Lepi- 
dolopha with a similar virgate shrubby habit is clearly a 
related genus. 

The list of species is compiled from Flora URSS (Komarov, 
1961) with one radiate species transferred from Pyrethrum by 
Muradyan (1970) added. The position of this species, X. 
tianshanicum, within Xylanthemum is provisional. According 
to Tzvelev it is more closely related to Trichanthemis , and 
differs from that genus simply by its glabrous cypselas. 
Tzvelev placed it in the monotypic Pyrethrum section 
Trichanthemopsis. Muradyan, when investigating cypsela 
morphology, transferred it to Xylanthemum mainly because 
of the presence of myxogenic cells on the cypselas. In other 
characters, e.g. of pappus and habit, the species appears 
more close to Trichanthemis or Richteria. Podlech in Podlech 
et al. (1986) considers this genus as a section of Tanacetum. 

X. fisherae (Aitch. & Hemsley) Tzvelev 

X. gilletii (Podl.) Bremer & Humphries, comb. nov. 

Basionym: Tanacetum gilletii Podl., Fl. iranica 158: 140 

(1986). 



X. lingulatum (Boiss.) Bremer & Humphries, comb. nov. 

Basionym: Pyrethrum lingulatum Boiss., Fl. orient. 3: 357 

(1875). 
X. macropodum (Hemsley & Lace) Bremer & Humphries, 

comb. nov. Basionym: Tanacetum macropodum Hemsley 

& Lace in J. Linn. Soc. 28: 324 (1891). 
X. paghmanense (Podl.) Bremer & Humphries, comb. nov. 

Basionym: Tanacetum paghmanense Podl., Fl. iranica 158: 

141 (1986). 
X. pamiricum (Hoffm.) Tzvelev 
X. polycladum (Rech. f.) Tzvelev 
X. rupestre (Popov ex Nevski) Tzvelev 
X. tianshanicum (H. Kraschen.) Muradyan (Pyrethrum tians- 
hanicum H. Kraschen.) 

19. LEPIDOLOPHA Winkler in Trudy imp. S. Peterb. 
hot. Sada 13: 236 (1894). Type species: L. 
komarovii Winkler. 

Shrublets with virgate, basally sometimes much woody stems. 
Leaves alternate or basally on the woody stems also fascicu- 
late on brachyblasts, entire or 3-lobed. Capitula solitary or 
corymbose, discoid. Involucre rather narrowly urceolate; 
involucral bracts in 4-7 rows, the outer much smaller than the 
inner. Receptacle flat to convex, epaleate. Corolla 5-lobed; 
tube basally swollen in fruit. Cypselas oblong, 6-10-ribbed, 
with sessile glands. Pappus of many subulate scales. 

Distribution. C. Asia. - 9 spp. 

Lepidolopha is well characterized, for example by the entire 
to 3-lobed leaves and the narrowly urceolate capitula with 
involucral bracts in several rows. The species are closely 
related with vicarious distributions and they differ mainly in 
foliage and inflorescence characters. A possibly related genus 
is Xylanthemum, also shrubby and with involucral bracts in 
several rows, though with a wider involucre and different 
pappus. 

Apart from the treatment by Knorring in Flora URSS 
(Komarov, 1961), there is also a more detailed discussion by 
the same author (Knorring, 1959). 

L. fedtschenkoana Knorr. 

*L. filifolia Pavlov 

*L. gomolitzkii Kovalevsk. & Safralieva 

L. karatavica Pavlov 

*L. komarovii Winkler {Tanacetum komarovii (Winkler) 

Muradyan) 
L. kraschenennikovii Kovalevsk. & Safralieva 
L. mogoltavica (H. Kraschen.) H. Kraschen. 
L. nuratavica H. Kraschen. {Tanacetum nuratavicum (H. 

Kraschen.) Muradyan) 
*L. talasica Kovalevsk. & Safralieva 

20. HIPPOLYTIA Polj. in Bot. Mater. Gerb. bot. 
Inst. V. A. Komarova 18: 288 (1957). Type species: 
H. darvasica (Winkler) Polj. 

Pubescent, sometimes densely tomentose perennial herbs. 
Leaves alternate and basally more or less rosulate, much 
pinnatisect. Capitula densely corymbose to glomerulate or 
rarely solitary, discoid. Involucral bracts with dark brown 
margins. Receptacle flat to convex, epaleate. Corolla 
5-lobed. Anthers tailed. Cypselas 5-8-ribbed, sometimes with 



106 

resin canals, with sessile glands, with a more or less distinct 
apical rim. Pappus absent. 

Distribution. C. Asia, Mongolia, China in Xinjiang and 
Tibet, and Himalaya (map by Shih, 1979: 68). - 19 spp. 

Poljakov (1957) discussed the wide circumscription and loose 
definition of Tanacetum (then including also Ajania) and he 
found it difficult to accept both homogamous (discoid) and 
heterogamous (disciform) species in the same genus. Conse- 
quently Poljakov removed a number of central Asian discoid 
Tanacetum species to a new genus, Hippolytia. The cypselas 
have no pappus as opposed to the Tanacetum species. The 
species have a characteristic habit with much pinnatisect 
leaves, pedunculoid stems with crowded (rarely solitary) 
capitula, and involucral bracts with dark brown margins. 
Some high-altitude species have densely tomentose leaves 
crowded on vegetative shoots and short flowering stems. This 
alpine habit is best explained as a secondary development 
within the genus. 

Tzvelev in Flora URSS (Komarov, 1961) stated that Hip- 
polytia is related to Dendranthema and Ajania and that 
Hippolytia is intermediate between these genera, a view 
analogous to the position of 'Cancrinia' (s. 1. incl. Tanacetop- 
sis) between Pyrethrum and Tanacetum s. s. According to 
Tzvelev radiate species of Dendranthema (cf. Pyrethrum) 
evolved into discoid species of Hippolytia (cf. Tanacetopsis) , 
which further evolved into disciform species of Ajania (cf. 
Tanacetum s. s.). 

Podlech et al. (1986) included Hippolytia as a section of 
Tanacetum. In our opinion Hippolytia, probably together 
with Heliocauta, is related to part of Tanacetum as discussed 
under Heliocauta. Hippolytia is retained here pending a 
reinvestigation of the whole Tanacetum complex. It is pos- 
sible that there are species of Ajania that should be trans- 
ferred to Hippolytia. 

Tzvelev removed some of Poljakov's species of Hippolytia 
to Ajania. Shih (1979) revised the genus and added a number 
of Chinese species. The list of species is taken from Shih's 
paper but with H. alashanensis listed as a synonym. 

H. crassicollum (Rech. f.) Bremer & Humphries, comb. nov. 
Basionym: Chrysanthemum crassicollum Rech. f. in Biol. 
Skr. 8 (2): 43 (1955) {Tanacetum crassicollum (Rech. f.) 
Podl.) 

H. darvasica (Winkler) Polj. 

H. delavayi (W. Smith) Shih 

H. desmantha Shih 

H. dolichophylla (Kitam.) Bremer & Humphries, comb. nov. 
Basionym: Chrysanthemum dolichophyllum Kitam. in 
Acta, phytotax. geobot. Kyoto 23: 73 (1968) (Tanacetum 
dolichophyllum (Kitam.) Kitam.) 

*H. glomerata Shih 

H. gossypina (C. B. Clarke) Shih 

H. herderi (Regel & Schmalh.) Polj. 

H. kaschgarica (H. Kraschen.) Polj. (//. alashanensis (Ling) 
Shih) - Note: In recent treatments of Hippolytia (Shih, 
1979) H. kaschgarica has disappeared. It was based on 
Tanacetum kaschgaricum H. Kraschen. (1933) from Tien- 
Shan range. Mount Ischma (China in Xinjiang). Material 
of this species in St. Petersburg, annotated by Kraschenin- 
nikov, appears to be the species currently known as Hip- 
polytia alashanensis (Ling) Shih originally described as 
Tanacetum alashanense Ling from Ala-Shan, Inner Mongo- 
Ha (China). At least, they must be closely related with the 



K. BREMER AND C. J. HUMPHRIES 

shrubby, subspinose habit (see Ling, 1983: plate 13,1). 
Furthermore, H. kaschgarica which is equal to H. alashan- 
ensis is aberrant in Hippolytia and thus deserves further 
study. 

*H. kennedyi (Dunn) Ling 

H. longifolia (Wallich) Shih (Tanacetum himachalense Aswal 
& Mehrotra) 

H. megacephala (Rupr.) Polj. 

*H. nana (C. B. Clarke) Shih 

H. schugnanica (Winkler) Polj. 

*H. senecionis (Besser) Polj. 

*//. syncalathiformis Shih 

H. tomentosa (DC.) Tzvelev 

*H. trifida (Turcz.) Polj. 

H. yunnanensis (Jeffrey) Shih 

2L HELIOCAUTA Humphries in Bot. Notiser 130: 
155 (1977). Type species: H. atlantica (Litard. & 
Maire) Humphries. 

A perennial rosulate or creeping herb. Leaves in a basal 
rosette, pinnatisect. Capitula solitary, pedunculate, discoid. 
Involucral bracts with dark brown margins. Receptacle coni- 
cal, paleate. Corolla 5-lobed; lobes soon brownish. Style 
immersed in a lobed nectary. Cypselas somewhat dorsiven- 
trally flattened, 4-5-ribbed with 2 major lateral ribs, with 
scattered elongated resin sacs and with an apical erose rim. 
Pappus absent. 

Distribution. N. Africa in Morocco. - Monotypic. 

This recently described monotypic genus is difficult to place 
and as stated by Humphries (1977) it is not related to 
Anacyclus, wherein it was formerly classified. Humphries 
compared Heliocauta to a number of genera, notably Achil- 
lea, but refrained from indicating a possible sister group. The 
Achillea species discussed by Humphries, A. barrelieri and A. 
oxyloba, are possibly plesiomorphic within the genus and 
within the subtribe Achilleinae. The similarities to Heliocauta 
may be interpreted as symplesiomorphies, shared also by 
members of Tanacetum. 

Another genus mentioned by Humphries is Sclerorhachis, 
supposed to have the same type of scattered epicarpic resin 
sacs. The elongated resin sacs in Heliocauta have transverse 
walls and are morphologically similar to the elongated rows 
of epicarpic myxogenic cells present in many genera. In 
Sclerorhachis, however, they are not resiniferous but myxo- 
genic as in other genera, and a close relationship between 
Heliocauta and Sclerorhachis cannot be assumed. 

Hippolytia is another possible relative of Heliocauta. They 
share a number of albeit homoplasious characters and can be 
placed together within 'Tanacetinae'. The reason for this 
placement is the presence of rather similar species within 
Tanacetum. It appears that both genera, singly or together, 
have their sister group within Tanacetum. One interesting 
species is T. tatsienense, a radiate species with short rays but 
otherwise very similar to Heliocauta and the single-headed 
species of Hippolytia. 

4. GONOSPERMINAE Bremer & Humphries, 
subtrib. nov. 

Type species: Gonospermum fruticosum (C. Smith ex Link) 
Less. 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



107 



Frutices vel herbae perennes, basi fruticosae. Folia multilo- 
bata lobis multis rotundatis vel interdum paucilobata vel 
Integra et margine dentata. Capitula laxe vel dense corym- 
bosa. Receptaculum paleaceum vel raro epaleaceum. Cypse- 
lae 5-10(-12)-costatae. Pappus e squamis parvis formatus, 
squama quoque costam terminanti. 

Shrubs or perennial, basally woody herbs. Leaves alternate, 
with many rounded lobes, sometimes few-lobed or entire and 
dentate. Capitula laxly to densely corymbose, radiate or 
discoid. Receptacle flat or conical to elongated, paleate or 
rarely epaleate. Ray floret limb white. Disc corolla 5-lobed. 
Cypselas 5-10(-12)-ribbed. Pappus of small scales, each 
terminating a rib. 

Distribution (Table 11). Canary Islands and southern 
Africa, mainly in Natal, one species also in Madagascar. - 3 
genera, 15 spp. 

Table 11 General distribution of Gonosperminae and Handeliinae 
and genera. x=indigenous, o=introduced. 



Table 12 Data matrix for the Gonosperminae. 1 = presence, = 
absence, ? = missing data or not applicable, p = polymorphic but 
scored as the plesiomorphic condition, a = polymorphic but 
scored as the apomorphic condition. 



C.&E. SW 
Asia Asia 



N.Afr. S.Afr. 



Gonosperminae 
Lugoa 

Gonospermum 
Inulanthera 






X 
X 
X 


X 
X 


Handeliinae 


X 


X 






Lepidolopsis 
Polychrysum 
Pseudohandelia 


X 

X 
X 


X 
X 
X 






Handelia 


X 


X 






Sclerorhachis 


X 









The interesting close relationship between the two Macaron- 
esian genera Lugoa and Gonospermum and the southern 
African Inulanthera has only recently been revealed 
(Kallersjo, 1986). Furthermore, the southern African group 
of species were formerly erroneously classified in Athanasia, 
obscuring their true relationships. The Gonosperminae is 
probably related to a part of Tanacetum, in the subtribe 
'Tanacetinae'. Of the two Macaronesian Tanacetum species, 
T. ferulaceum and T. ptarmiciflorum , the latter at least is a 
possible close relative. 

Clades and characters - Fig. 5, Tables 2, 12. 

\=GolTf= 22 Lugoa 

rj= 23 Gonospermum 
ll=Go2iJ= 24 Inulanthera 

Fig. 5 Cladogram of the Gonosperminae produced by the ie option 
in Hennig86. Cladogram length = 9, consistency index = 100, 
retention index = 100. 

Clade Gol - subtribe Gonosperminae 

19 Leaves large with many rounded lobes. Inulanthera is very 
variable in leaf shape, but there are also species of Inulan- 
thera with these kinds of leaves, very similar to those of 
Gonospermum and Lugoa. 

45 Receptacle paleate. One species of Inulanthera is epaleate. 
162 Pappus of scales or teeth projected from the ribs. 



nil 1 1 

1450576 146 23992 

5327181 952295254 



11 11 

177555556666617 
356356790567851 



22. Lugoa alaal?l lalOOOOOO ?? 7000000000000 

23. Gonospermum al?al?l lallllOOO ??????????????? 

24. Inulanthera al?al?l aalllllll ??????????????? 



Lugoa 

There is no obvious autapomorphy for this genus. 

Clade Go2 

2 Plants shrubby. 

29 Capitula densely corymbose. 

35 Capitula discoid. 

Gonospermum 

There is no obvious autapomorphy for this genus, which 
appears undefined by comparison to Inulanthera. 

Inulanthera 

92 Anthers caudate. 

95 Anthers with endothecial tissue partly or wholly polarized. 

124 Cypselas with 10 (8-12) multicellular epicarpic ribs. 

22. LUGOA DC, Prodr. 6: 14 (1838). Type species: 
L. revoluta (C. Smith ex Link) DC. 

A suffruticose, basally woody perennial. Leaves alternate, 
pinnatisect, large and flat with rounded lobes. Capitula laxly 
corymbose, radiate. Receptacle conical, paleate. Ray florets 
female, fertile; limb white. Disc corolla 5-lobed, shghtly 
thickened at base. Cypselas 5-ribbed. Pappus of teeth pro- 
jected from the ribs. 

Distribution. Canary Islands. - Monotypic. 

Lugoa with radiate capitula is the most plesiomorphic mem- 
ber of subtribe Gonosperminae. There is no obvious autapo- 
morphy for this monotypic genus, though the one species is 
clearly distinct from the species of Gonospermum. It is 
sometimes included in Gonospermum, but retained here in 
accordance with recent practice (e.g. Bramwell &. Bramwell, 
1974). Furthermore, inclusion of Lugoa in Gonospermum 
would result in a clearly paraphyletic taxon since the discoid 
Gonospermum species are more closely related to Inulan- 
thera than to Lugoa. 

23. GONOSPERMUM Less., Syn. gen. Compos.: 263 
(1832). Selected type species: G. fruticosum (C. 
Smith ex Link) Less. 

Rather elaborate shrubs. Leaves alternate, pinnatisect, large 
and flat with many rounded lobes. Capitula corymbose, 
discoid. Receptacle elongated, paleate. Corolla 5-lobed, 



108 



K. BREMER AND C. J. HUMPHRIES 



slightly thickened at base. Cypselas 5-ribbed. Pappus of 
scales or teeth projected from the ribs. 

Distribution. Canary Islands. - 4 spp. 

Gonospermum is related to Inulanthera and Lugoa. Inulan- 
thera differs by its caudate anthers and many-ribbed cypselas. 
By comparison to Inulanthera, Gonospermum is undefined 
and paraphyletic. Bramwell & Bramwell's (1974) Wild flow- 
ers of the Canary Islands is useful for species identification. 

G. canariense (DC.) Less. 

G. elegans (Cass.) DC. 

G. fruticosum (C. Smith ex Link) Less. 

G. gomerae Bolle 

24. INULANTHERA Kallersjo in Nord. J. Bot. 5: 539 
(1986). Type species: /. calva (Hutch.) Kallersjo. 

Shrubs or rarely basally woody half-shrubs. Leaves alternate, 
variously lobed, dentate or entire. Capitula corymbose, dis- 
coid. Receptacle flat, paleate, rarely epaleate. Corolla 
5-lobed, with a narrow tube and a distinct limb. Anthers 
caudate; endothecial tissue polarized. Cypselas 8-10-ribbed, 
glabrous or sometimes glandular. Pappus of small scales or 
teeth, each terminating a rib. 

Distribution. S. Africa in the E. Cape, Natal, and Trans- 
vaal, and in Lesotho, Angola (/. schistostephioides) , Zimba- 
bwe (/. nuda) and Madagascar (/. brownii). - 10 spp. 

Inulanthera is recently described and is comprised of a 
number of species formerly included in Athanasia. The 
species of Inulanthera were found to differ considerably from 
Athanasia in cypsela anatomy, having sclerenchyma of longi- 
tudinal bundles or elongated cells in the ribs rather than 
sclerified parenchyma cells as in Athanasia and lacking resin 
canals which are present in Athanasia. The species of Inulan- 
thera investigated chemically also have the common poly- 
acetylenes rather than furanosesquiterpenes, which 
characterize Athanasia and the group of related genera within 
the Ursiniinae. In these features Inulanthera is similar to most 
Anthemideae but the genus is further characterized by its 
basally caudate anthers, a very unusual character in 
Anthemideae (occurring elsewhere only in Hippolytia and 
Osmitopsis). 

Cypselas with ribs projected into small pappus scales or 
teeth, and the characteristic leaves with many rounded leaf 
lobes of some species (/. brownii, I. nuda, I. schistostephio- 
ides) clearly indicate a relationship to Gonospermum and 
Lugoa. Other species have few-lobed or dentate leaves, 
presumably apomorphic conditions within the genus. The 
South African species, all occurring in Natal, are at the 
species level adequately described under Athanasia by Mill- 
iard (1977). 

/. brownii (Hochr.) Kallersjo {Athanasia brownii Hochr.) - 
Note: Description in Hochreutiner, 1908. 

/. calva (Hutch.) Kallersjo {Athanasia calva Hutch.) 

/. coronopifolia (Harvey) Kallersjo {Athanasia coronopifolia 
Harvey) 

/. dregeana (DC.) Kallersjo {Athanasia dregeana (DC.) Har- 
vey, Athanasia punctata (DC.) Harvey) 

/. leucoclada (DC.) Kallersjo {Athanasia leucoclada (DC.) 
Harvey) 

/. montana (J. M. Wood & M. Evans) Kallersjo {Athanasia 
montana J. M. Wood & M. Evans) 



/. nuda Kallersjo {Pentzia schistostephioides M. Taylor) - 
Note: Description in Taylor, 1940. 

/. schistostephioides (Hiern) Kallersjo {Athanasia schis- 
tostephioides Hiern) - Note: Description in Hiern, 1898. 

/. thodei (Bolus) Kallersjo {Athanasia thodei Bolus) 

/. tridens (Oliver) Kallersjo {Athanasia tridens Oliver) 

5. HANDELIINAE Bremer & Humphries, 
subtrib. nov. 

Type species: Handelia trichophylla (Schrenk) Heimerl. 

Herbae perennes, basaliter suffruticosae et villosae- 
tomentosae, caule baud vel pauceramoso crassiusculo 
medulla moUi. Folia alterna vel rosulata, pinnatisecta lobis 
filiformibus. Capitula dense vel laxe corymbosa, raro anguste 
paniculata, discoidea. Cypselae plerumque parvae, quinque- 
costatae. Pappus coroniformis e squamis parvis formatus vel 
nuUus. 

Basally more or less woody and villous-tomentose perennials 
with generally few- or unbranched rather thick stems with a 
soft pith. Leaves alternate or rosulate, pinnatisect with fiH- 
form lobes. Capitula generally small in dense to lax corymbs 
or rarely in a long narrow panicle, discoid. Receptacle flat to 
conical, paleate or epaleate. Corolla 5-lobed. Cypselas gener- 
ally small and 5-ribbed, rarely slender with obtuse excres- 
cences, sometimes with myxogenic cells. Pappus a corona of 
small scales or absent. 

Distribution (Table 11). Asia, mainly central part. - 5 
genera, 8 spp. 

This subtribe forms a homogeneous and monophyletic group, 
characterized by a number of synapomorphies. The relation- 
ship of Lepidolopsis , Poly chry sum, Handelia and Pseudohan- 
delia was first noted by Tzvelev (in Komarov, 1961). 
Sclerorhachis was originally part of Anthemis, mainly because 
of its paleate receptacle but after it was placed as a section of 
Anthemis (Rechinger, 1944) Rechinger elevated it to generic 
rank of uncertain position within the tribe (Rechinger 1968). 
We consider that Sclerorhachis is related to Handelia and 
Pseudohandelia and a true member of this subtribe. This 
relationship has probably been obscured by the distinct habit 
of Sclerorhachis, with basal leaf- rosettes and laxly branched, 
leafless aerial stems. We interpret these stems as branches of 
a secondarily lax inflorescence, homologous to the dense 
corymbs of the other genera. 

From the data matrix there is only one parsimonious 
solution to the generic interrelationships within Handeliinae, 
shown in the cladogram. The sister group of Handeliinae is 
probably to be found within Tanacetum, where there are 
several thick-stemmed species with corymbose inflores- 
cences. 

Clades and characters - Fig. 6, Tables 2, 13. 



^Hal 



E 



25 Lepidolopsis 

!(= 26 Polychrysum 
Ha2l [r" 27 Pseudohandelia 

0-:Ha3| rp= 28 Handelia 

li=Ha4:li=^ 29 Sclerorhachis 



Fig. 6 Cladogram of the Handeliinae produced by the ie option in 
Hennig86. Cladogram length = 13, consistency index = 92, 
retention index = 80. 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 

Table 13 Data matrix for the Handeliinae. 1 = presence, = 
absence, ? = missing data or not applicable, p = polymorphic but 
scored as the plesiomorphic condition, a = polymorphic but scored 
as the apomorphic condition. 

1111 111 nil 11 

1450576 13323701457 1555556666617775674 
5327181 865190293511 3356790567851560606 

25. Lepidolopsis al?al?l 111100000000 linilllllllllllOQp 

26. Poly chry sum al?al?l 111011000000 ????????????????000 

21 .Pseudohandeliaa\la\l\ 111010111000 111111111111111111^ 

2%.Handelia al?al?l lllOlOlOOaOO lllllllllllllllOllp 

29. Sclerorhachis al?al?l lalOOOlOOaal 1111111111111110110 



Clade Hal - subtribe Handeliinae 

8 Plants basally villous-tomentose with rather thick stems and a 
soft pith. 

16 Leaves much pinnatisect with filiform lobes. 

35 Capitula discoid. 

Lepidolopsis 

31 Capitula in a long narrow panicle or raceme. 

Clade Ha2 

29 Capitula densely corymbose. In Sclerorhachis the capitula 
are terminal on the loosely branched, leafless aerial stems, 
which are considered homologous to the branches of a 
corymbose inflorescence. 

Polychrysum 

30 Capitula very small and numerous in a large, dense, 
semiglobose corymb. 

Clade Ha3 

172 Pappus absent in ray and disc cypselas. 

Pseudohandelia 

109 Cypselas arcuate. 

113 Cypselas slender and tuberculate with numerous obtuse 
excrescences. 

Clade Ha4 

45 Receptacle paleate. 

Handelia 

There is no obvious autapomorphy for this genus. 

Sclerorhachis 

29 reversed. See clade Hal. 

51 Floral parts with resin canals. 

71 Corolla apically contracted. 

25. LEPIDOLOPSIS Polj. in Bot. Mater. Gerb. hot. 
Inst. V. A. Komarova 19: 374 (1956). Type species: 
L. turkestanica (Regel & Schmalh.) Polj. 

A basally woody perennial with rather thick unbranched 
stems with a soft pith, villous-tomentose at base and the leaf 
bases. Leaves alternate, much pinnatisect with filiform lobes. 
Capitula small, numerous in a long panicle, discoid. Recep- 
tacle conical, epaleate. Corolla 5-lobed, with sessile glands. 



109 

Cypselas small, 5-ribbed. Pappus of several mainly abaxial 
acute scales. 

Distribution. Iran, Afghanistan and C. Asia. - Monotypic. 

Vegetatively Lepidolopsis is very similar to Polychrysum, 
Handelia and Pseudohandelia. All four genera have single, 
unbranched or apically few-branched, basally pubescent 
stems with much pinnatisect leaves with filiform lobes. Lepi- 
dolopsis differs by its much elongated inflorescence and its 
sessile floret glands. Originally Poljakov included a number 
of discoid species in Lepidolopsis. These are now in other 
genera, mainly Tanacetum and Tanacetopsis . Both Poly- 
chrysum tadshikorum and Pseudohandelia umbellifera were 
also classified in Lepidolopsis by Poljakov. 

26. POLYCHRYSUM (Tzvelev) Kovalevsk. in 
Vvedensky, Fl. uzbekistana 6: 148 (1962). Type 
species: P. tadshikorum (Kudr.) Kovalevsk. - 
Cancrinia sect. Polychrysum Tzvelev. 

A basally woody perennial, one to few-branched with rather 
thick light stems and a soft pith, villous-tomentose at the leaf 
bases. Leaves alternate, much pinnatisect with filiform lobes. 
Capitula small, numerous in a large, dense, semiglobose 
corymb, discoid. Receptacle convex, epaleate. Corolla 
5-lobed, with stalked glands. Cypselas small, 5-ribbed, with 
stalked glands. Pappus a corona of many small scales. 

Distribution. Afghanistan, C. Asia. - Monotypic. 

Polychrysum is based on a monotypic section of Cancrinia, 
described by Tzvelev in Flora URSS (Komarov, 1961). It is 
clearly different from the type species of Cancrinia, and more 
closely related to Lepidolopsis , Handelia and Pseudohande- 
lia. One character stressed by Tzvelev is the capitulum with 
only one mature fruit. This is obviously correlated to the 
small and numerous capitula. 

27. PSEUDOHANDELIA Tzvelev in Komarov, Flora 
URSS 26: 878 (1961). Type species: P. umbellifera 
(Boiss.) Tzvelev. 

A basally woody perennial with unbranched, rather thick 
stems with a soft pith, villous-tomentose at the stem and leaf 
bases. Leaves alternate, basally approaching rosulate, much 
pinnatisect with filiform lobes. Capitula corymbose, discoid. 
Receptacle conical, epaleate. Corolla 5-lobed, with many 
stalked glands. Cypselas slender and somewhat arcuate, with 
5 vascular strands, tuberculate with numerous obtuse excres- 
cences. Pappus absent. 

Distribution. Iran, Afghanistan, C. Asia and China in 
Xinjiang. - Monotypic. 

Pseudohandelia and Handelia are very similar and closely 
related. Their close relationship has been more or less 
concealed by the traditional practice of keeping taxa with and 
without receptacular paleae far apart in classification. The 
two genera are often confused. Apart from the receptacle 
character Pseudohandelia is also distinguished by its slender 
and tuberculate cypselas and the more dense, umbelliform 
corymb. The close relationship of Pseudohandelia to Hande- 
lia as well as to Polychrysum (treated by Tzvelev as a section 
of Cancrinia), and Lepidolopsis was first noted by Tzvelev (in 
Komarov, 1961). 



no 



K. BREMER AND C. J. HUMPHRIES 



The cypsela excrescences are homologous with gland 
stalks; glands with such stalks are present on the corolla. 



28. HANDELIA Heimerl in Ost. bot. Z. 71: 215 
(1922). Type species: H. trichophylla (Schrenk) 
Heimerl. 

A basally woody perennial with few- or unbranched, rather 
thick stems with a soft pith, basally and at the leaf bases 
villous-tomentose. Leaves alternate, basally becoming almost 
rosulate, leaves pinnatisect, with deep sinuses and filiform 
lobes. Capitula small, numerous, corymbose, discoid. Recep- 
tacle conical, paleate. Corolla 5-lobed, with stalked glands. 
Cypselas small, 5-ribbed, with an abaxial rim. Pappus absent. 

Distribution. Afghanistan, Pakistan, C. Asia and China in 
Xinjiang. - Monotypic. 

Handelia is very similar to and closely related to Pseudohan- 
delia, though the two genera have been placed far apart. 
Handelia has a paleate receptacle, whereas Pseudohandelia is 
epaleate. They are often confused but they are easily distin- 
guished by the receptacle character and by their different 
cypselas. Handelia also has a more laxly branched inflores- 
cence than Pseudohandelia. The relationship to Sclerorhachis 
is discussed below. 



29. SCLERORHACHIS (Rech. f.) Rech. f. inAnz. 
ost. Akad. Wiss. Mathematische 
Naturwissenschaftliche Klasse 105: 242 (1968). Type 
species: 5. caulescens (Aitch. & Hemsley) Rech. f. - 
Anthemis sect. Sclerorhachis Rech. f. 

Rosulate basally pubescent perennials with a basal leaf- 
rosette and almost naked, loosely branched stems, basally 
rather thick and with a soft pith. Leaves pinnatisect, with a 
persistent, sclerified rachis. Capitula laxly corymbose, pedun- 
culate, discoid. Receptacle flat to convex, paleate; paleae 
filiform. Corolla 5-lobed, with a distinct, apically contracted 
limb. Cypselas 5-ribbed, with scattered rows of myxogenic 
cells. Pappus absent. 

Distribution. Iran and Afghanistan. - 4 spp. 

The type species of Sclerorhachis was originally placed in 
Anthemis, though considered highly isolated (Rechinger, 
1944). At a first glance Sclerorhachis seems isolated but it is a 
member of subtribe Handeliinae together with Handelia and 
Pseudohandelia (Rechinger, 1955, 1968). In Sclerorhachis the 
leaves are concentrated into a basal leaf-rosette and we 
consider the loosely branched, leafless aerial stems with 
terminal capitula homologous to the corymbose inflorescence 
of the other genera. The receptacle is furnished with filiform 
bristles in the same position as receptacular paleae, hence 
they appear homologous to paleae. 

S. caulescens (Aitch. & Hemsley) Rech. f. {Anthemis caule- 
scens Aitch. & Hemsley) 
*S. leptoclada Rech. f. 
5. platyrhachis (Boiss.) Podl. ex Rech. f. 
S. polysphaera Rech. f. 



6. ARTEMISIINAE Less, emend. Bremer & 
Humphries, emend, nov. 

Lessing in Linnaea 5: 163 (1830) ('Artemisieae'). Type spe- 
cies: Artemisia vulgaris L. 

Herbae annuae vel perennes, suffrutices vel frutices. Capitula 
interdum radiata, saepe disciformia vel discoidea, corymbosa 
vel paniculata. Receptaculum plerumque epaleaceum. 
Appendix apicalis antherarum subtriangularis vel lanceolato- 
linearis, parietibus cellularum aliquantum incrassatis. Cypse- 
lae oblongo-obovoideae, quinque-costatae vel saepe 
obovoideae ecostatae, parietibus tenuibus, raro pilosae. Pap- 
pus nullus vel raro coroniformis e squamis parvis formatus. 

Perennial, rarely annual herbs, half-shrubs or shrubs; indu- 
mentum frequently of dolabriform hairs. Leaves variously 
dissected, rarely entire. Capitula radiate or generally disci- 
form or discoid, often rather small; inflorescence various but 
often corymbose or paniculate. Receptacle epaleate or very 
rarely paleate. Ray floret limb white, yellowish or pink. 
Outer florets (in disciform capitula) in one row (rarely two), 
female, fertile. Disc or central florets perfect and hermaphro- 
dite or female-sterile, 5-lobed. Apical anther appendages 
(sub)triangular-lanceolate-linear, of rather thick-walled cells. 
Pollen often without or with short spines. Cypselas oblong- 
obovoid, 5-ribbed or generally rather small, obovoid and 
faintly ribbed or without ribs, thin-walled, with or without 
myxogenic cells in rows, rarely pilose. Pappus absent or very 
rarely coroniform, of small scales. 

Distribution (Table 14). Worldwide but mainly northern 
hemisphere and especially central and E. Asia, some Artemi- 
sia species widespread as weeds. - 18 genera, 634 spp. 

The main genera of this subtribe are familiarly known as the 
Artemisia group, including the large genera Artemisia and 
Seriphidium (commonly considered a section of Artemisia), 
eight small Asian genera, and two small North American 
genera. The group is characterized by disciform or discoid, 
commonly paniculate capitula, smooth or short spined pol- 
len, and obovoid, thin-walled, and ribless cypselas without a 
pappus. 

However, within this subtribe we have included the prob- 
able, more plesiomorphic relatives of the Artemisia group 
(Fig. 7, Clades Arl-Ar5), which in previous classifications 
were hidden within a broad concept of Chrysanthemum. 
These relatives include the radiate genera Brachanthemum, 
Dendranthema, Arctanthemum and Tridactylina with solitary 
or laxly corymbose capitula, and the disciform Ajania and 
Phaeostigma with densely corymbose capitula. These genera 
have rather thin-walled and more or less faintly ribbed 
cypselas always without a pappus. The apical anther append- 
ages are synapomorphic for the whole subtribe. They are 
(sub)triangular to lanceolate-linear, often acute to acumi- 
nate, and composed of rather thick-walled cells, best 
expressed in the apomorphic representatives of the Artemisia 
group. 

The sister group of Artemisiinae is to be found within 
Tanacetum, where there are species very similar to represen- 
tatives of Dendranthema and Brachanthemum. The delimita- 
tion between Tanacetum and these two genera, notably 
Dendranthema, traditionally rests on two characters, pres- 
ence of pappus and myxogenic cells on the cypselas. Pappose 
species without myxogenic cells are classified in Tanacetum 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 

Table 14 General distribution of Artemisiinae and genera. x=indigenous, o=introduced. 



Ill 



N. Am. Eur- 
Asia 



C. & E. SW 

Asia Asia 



S. Eur. N. Afr. S. Afr. Austr. S. Am. 
N. Zeal. 



Artemisiinae 


X 


X 


X 


X 


X 


Brachanthemum 






X 






Dendranthema 




X 


X 






Arctanthemum 




X 


X 


X 




Tridactylina 






X 






Ajania 






X 






Phaeostigma 






X 






Stilpnolepis 






X 






Ajaniopsis 






X 






Filifolium 






X 






Sphaeromeria 


X 










Kaschgaria 






X 






Seriphidium 


X 


X 


X 


X 


X 


Crossostephium 








X 




Artemisia 


X 


X 


X 


X 


X 


Neopallasia 






X 






Turaniphytum 






X 






Mausolea 






X 






Picrothamnus 


X 











and epappose species with or without myxogenic cells are 
classified in Dendranthema. (Pappose species with myxogenic 
cells represent another problem, and are more or less provi- 
sionally relegated to other genera, as exemplified by Xylan- 
themum tianshanicum) . The relationship of Dendranthema 
and Artemisiinae in general to Tanacetum cannot be assessed 
in detail before the latter genus has been properly revised and 
dismantled. 

The cladogram (Fig. 7) is one of many (71) equally 
parsimonious trees. In this case the analysis was performed 
with characters 29 and 31 combined into one multistate 
character with two apomorphic states, densely corymbose 
and paniculate-racemose capitula. A preliminary analysis 
with the binary characters as presented in the matrix (Table 
15) yielded an artificial grouping of Stilpnolepis and 
Seriphidium supported by a reversal in character 29, i.e. loss 
of the dense capitula corymbs, but the two genera are totally 
different in inflorescence structure. The strict consensus tree 
of all the equally parsimonious cladograms is almost totally 
collapsed, indicating the instability in our current hypothesis 
of Artemisiinae generic interrelationships. However, clade 
Ar2 with Dendranthema, Arctanthemum and Tridactylina, as 
well as clade ArlO, Mausolea and Picrothamnus, were sup- 
ported by all cladograms and hence retained in the consensus 
tree. 

Clades and characters - Fig. 7, Tables 2, 15. 

Clade Arl - subtribe Artemisiinae 

11 Plants with dolabriform hairs. Dolabriform, i.e. T-shaped, 
or Y-shaped hairs occur in most genera of this subtribe. They 
are also frequent in Leucantheminae, Anthemis (Anthemidi- 
nae) and some genera of Achilleinae. The character is 
possibly a synapomorphy at a lower level within the tribe. 

93 Anthers with triangular-linear-lanceolate apical append- 
ages, of rather thick-walled cells. This character is present in 
all genera, though variously expressed. 

172 Pappus absent in ray and disc cypselas. Crossostephium 



and one species of Sphaeromeria both have a pappus of short 
scales. 

Brachanthemum 

2 Plants shrubby. In Artemisiinae there are many transitions 
from woody perennials to half-shrubs or shrubs. Hence the 
character is difficult to apply and it is present within several 
genera not scored in the matrix. 

Clade Ar2 

44 Involucral bracts with dark brown margins. This character 
is present also within Ajania, Phaeostigma, and Tanacetum, 
where the sister group of the subtribe is found. Hence the 
character may be plesiomorphic within the subtribe. 

Dendranthema 

This genus is plesiomorphic compared to Arctanthemum, 
Tridactylina and possibly also compared to the rest of the 
subtribe. 

Arctanthemum 

6 Plants rhizomatous with rosulate, spathulate-obovate-linear 
leaves. 

15 reversed. Leaves not variously deeply lobed or divided, but 
entire. 

Tridactylina 

1 Plants annual. 

52 reversed. Ray floret limb not white, but yellow. There are 
several species of Dendranthema with yellow rays. 

Clade Ar3 

29 Capitula densely corymbose. Inflorescences are variable 
and complicated in subtribe Artemisiinae. Most of the Arte- 
misia group of genera have paniculate inflorescences. In 
Stilpnolepis the capitula are, probably secondarily, solitary or 
laxly corymbose. 



112 



K. BREMER AND C. J. HUMPHRIES 



fF= 30 Brachanthemum 



[=Arl= 



h=Ar2 






31 Dendranthema 

32 Arctanthemum 

33 Tridactylina 
rpr 34 Ajania 

li=Ar3| n= 35 Phaeostigma 

IL=Ar4| n= 36 Stilpnolepis 



'L 



Ar6 



37 Ajaniopsis 

38 Filifolium 
[[= 39 Sphaeromeria 






% 



ArS 



[ 



40 Kaschgaria 

41 Seriphidium 

42 Crossostephium 
j= 43 Artemisia 



Ar9:^ 



[ 



44 Neopallasia 

45 Turaniphytum 

r= 46 Mausolea 



^ArlO:^ 47 Picrothamnus 

Fig. 7 Cladogram (of 71 possible) of the Artemisiinae produced by the bb option in Hennig86. Cladogram length = 48, consistency index = 
79, retention index = 82. 

Table 15 Data matrix for the Artemisiinae. 1 = presence, = absence, ? = missing data or not applicable, p = polymorphic but scored as the 
plesiomorphic condition, a = polymorphic but scored as the apomorphic condition. 



Ill 
145057 
532718 



1 11 1 1 111 

197 4 23905369741318312916936700466 8 
13224619662455783312086297612494687831 



1 1 11 11 11 

5677755556175531 147 46788 
06056357905161744657983223 



30. Brachanthemum a 1 a a 1 ? 

3 1 . Dendranthema a 1 a a 1 1 

32. Arctanthemum 01111? 

33. TridactyUna a 1 ? a 1 ? 

34. Ajania a 1 ? a 1 ? 

35. Phaeostigma a 1 ? a 1 ? 

36. Stilpnolepis a 1 ? a 1 ? 

37. Ajaniopsis a 1 ? a 1 ? 

38. Filifolium a 1 ? a 1 ? 

39. Sphaeromeria a 1 ? a 1 ? 

40. Kaschgaria a 1 ? a 1 ? 

41 . Seriphidium a 1 ? a 1 ? 

42. Crossostephium a 1 ? a 1 ? 

43. Artemisia a 1 ? a 1 1 

44. Neopallasia a 1 ? a 1 ? 

45. Turaniphytum a 1 ? a 1 ? 

46. Mausolea a 1 ? a 1 ? 

47. Picrothamnus a 1 ? a 1 ? 



al 

al 

al 

?1 

al 

al 

?1 

?1 

al 

al 

al 

al 

?10 1 

al 

al 

?1 



al 
al 



0000 
aOOp 
1100 
1010 
pOOa 
pOOl 
OOpO 
001 1 
0001 
000a 
0000 
OOpO 
0000 
pOpO 
OOaO 
0000 
0000 
0000 



0000 
0000 
0000 
0000 
lOpO 
1110 
0101 
1 101 
1101 
1101 
1101 
0101 
1 100 
1101 
1 101 
1 101 
1 101 
1101 



pOOOO 
00000 
00000 
00000 
00000 
00000 
l?p00 
01 1 10 
01 101 
01 100 
01 100 
17100 
01 100 
01 100 
01 100 
01 100 
01 100 
01 100 



?0p0000 
7000000 
7000000 
7000000 
7000000 
7000000 
7000000 
7000000 
7000000 
lOpOOOO 
71 1 1000 
laOOllO 
7100001 
aapOOOO 
7100000 
7100000 
7100000 
1 100000 



00000000 
00000000 
00000000 
00000000 
00000000 
00000000 
00700000 
00000000 
00000000 
00000000 
00000000 
00700000 
00000000 
lOOOOOOp 
11 110000 
10001 1 10 
10000001 
10000001 



000000 
000000 
000000 
000000 
000000 
000000 
007700 
000000 
000000 
OOOOOp 
000000 
007700 
000000 
pOOOpO 
000000 
000000 
111100 
1 1001 1 



7 7 7 7 70 
7 7 7 7 70 
777000 
7 7 7 7 70 

9 9 9 9 9 9 



0000000 
0000000 
0000000 
0000000 

9 9 9 9 9 9 9 



7 7 7 7 7? 

9 9 9 9 9 9 



??????? 
9 9 9 9 9 9 9 



7 7 7 7 7 7 

9 9 9 9 9 9 



9 9 9 9 
9 9 9 9 



770 

9 9 9 



7 7 7 7 7 7 

9 9 9 9 9 9 



7 7 7 7 

9 9 9 9 



7 7 7 7 7 7 
700777 

9 9 9 9 9 9 



7 7 7 7 
9 9 9 9 



7 7 7 7 7? 

9 9 9 9 9 9 



7 7 7 7 

9 9 9 9 



7 7 7 7 7 7 

9 9 9 9 9 9 



9 9 9 9 
9 9 9 9 
9 9 9 9 



7 7 7 

9 9 9 

9 9 9 

9 9 9 

9 9 9 

9 9 9 

9 9 9 

9 9 9 

9 9 9 



00000 
00000 

00000 
00000 
ppOOO 
00000 
pOpOO 
00000 
00000 
OOOpO 
00000 
OpOOp 
00000 
OpOpp 
OOOOp 
00000 
00000 
00000 



0000 
0000 
0000 
0000 
0000 
0000 
0000 
0000 
0000 
0000 
0000 
ppOO 
0000 
OOpp 
0000 
0000 
0000 
0000 



0000 
0000 
0000 
0000 
0000 
0000 
0000 
0000 
0000 
0000 
0000 
0000 
000 

pppp 

0000 
0000 
0000 
0000 



36 Capitula disciform. Stilpnolepis and Seriphidium have 
discoid capitula. Within this subtribe discoid capitula are 
most parsimoniously interpreted as derived from disciform, 
rather than from radiate capitula. 

Ajania 

There is no autapomorphy for this genus, which appears to be 
a paraphyletic grade when compared to Phaeostigma and the 
Artemisia group. 

Clade Ar4 

96 Pollen grains with short spines or without spines. 

Phaeostigma 

102 Style-branches brownish. 

Clade Ar5 - the Artemisia group 

154 Cypselas thin-walled, obovoid to oblanceolate, devoid of 
ribs. Faint ribs are sometimes present, most clearly seen in 
Crossostephium and sometimes within Sphaeromeria. 

Stilpnolepis 



29 reversed. See clade Ar3. 

35 Capitula discoid. See character 36 under clade Ar3. 

Clade Ar6 

65 Outer female floret corollas 'flask-shaped' , tapering above 
or narrowly cylindrical. 

97 Pollen grains without spines. 

Ajaniopsis 

1 Plants annual. 

78 Corolla apically with erect, straight hairs. 

Filifolium 

143 Cypselas with myxogenic glands in 2 distinct adaxial- 
lateral rows. 

Clade Ar7 

13 Plants with interxylary cork. Interxylary cork has been 
studied in Sphaeromeria, Seriphidium, Artemisia and Picroth- 
amnus only. 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 

Sphaeromeria 

There is no obvious autapomorphy for this genus. 

Clade Ar8 

29 reversed. See clade Ar3. 

31 Capitula in a long narrow panicle or raceme. See character 
29 under clade Ar3. 

Kaschgaria 

12 Plants with stellate hairs. 

80 Corolla apically with stellate hairs. 

Seriphidium 

35 Capitula discoid. See character 36 under clade Ar3. 

38 Involucral bracts in 4-7 rows. 

116 Disc cypselas laterally flattened. 

Crossostephium 

2 Plants shrubby. See note under Brachanthemum. 

22 Leaves entire or apically tridentate. 

154 reversed. See clade Ar5. 

172 reversed. See clade Arl. 

Clade Ar9 

99 Style slender, parallel-sided at base. 

Artemisia 

There is no autapomorphy for Artemisia when compared to 
the remaining four genera. 

Neopallasia 

1 Plants annual. 

17 Leaves pectinate-pinnatisect with filiform, apically some- 
what swollen and mucronulate lobes. 

66 Outer female floret corollas without teeth. 

91 Central florets of two kinds; outer perfect, inner completely 
sterile with reduced ovaries. 

Turaniphytum 

32 Capitula in glomerules arranged in long spikes. 

64 Outer female florets subtended by scaphoid bracts. 
79 Corolla apically with long, reddish hairs. 
Clade ArlO 

2 Plants shrubby. See note under Brachanthemum. 

104 Disc floret style-branches fused. This is characteristic also 
of Artemisia sect. Dracunculus. 

106 Central floret ovaries reduced; florets functionally male. 

148 Cypselas cobwebby pilose. 

Mausolea 

67 Outer female florets without corollas. 

68 Outer female floret style-branches lanceolate, flat, acute. 
Picrothamnus 



113 
3 Plants spiny. 
81 Corolla cobwebby pilose. 

30. BRACHANTHEMUM DC, Prodr. 6: 44 (1838). 
Type species: B. fruticulosum (Ledeb.) DC. 

Small, more or less procumbent shrublets, woody at the base. 
Leaves alternate, few-lobed. Capitula solitary or laxly corym- 
bose, pedunculate, radiate or rarely discoid. Receptacle flat 
or convex to conical, epaleate. Ray florets female, fertile; 
limb yellow or yellowish. Disc corolla 5-lobed; tube with 
sessile glands. Apical anther appendage subtriangular. 
Cypselas obovoid to oblong, faintly 5-7-ribbed, thin-walled, 
with myxogenic cells. Pappus absent. 

Distribution. C. Asia, Mongolia and China (map by Kra- 
scheninnikov, 1949: 199). - 10 spp. 

According to Krascheninnikov (1949) and Tzvelev (in Koma- 
rov, 1961) most species of Brachanthemum are closely 
related, although B. baranovii is different and placed in the 
monotypic sect. Dendranthemopsis. It differs from sect. 
Brachanthemum by its oblong rays and flat, shortly pilose 
receptacle, as opposed to the otherwise shorter rays and 
convex to conical, glabrous receptacle. On the other hand the 
characteristic shrubby habit, the few-lobed leaves, and the 
rather small, cyathiform to urceolate capitula may distinguish 
this genus as monophyletic. Tzvelev noted the close relation- 
ship to Dendranthema, emphasized also by the similar thin- 
walled, myxogenic fruits without a pappus (described in 
detail by Savczenko, 1949). The list of species is taken from 
Krascheninnikov's (1949) revision and Flora URSS (Koma- 
rov, 1961) with species from China and Mongolia added. 

*B. baranovii (H. Kraschen. & Polj.) H. Kraschen. 

B. fruticulosum (Ledeb.) DC. 

B. gobicum H. Kraschen. 

B. kasakhorum H. Kraschen. 

B. kirghisorum H. Kraschen. 

*B. krylovii Serg. 

B. mongolicum H. Kraschen. 

*B. mongolorum Grubov 

B. pulvinatum (Hand.-Mazz.) Shih {B. nanshanicum H. 

Kraschen.) 
B. titovii H. Kraschen. 

31. DENDRANTHEMA (DC.) Des MouL in Act. Soc. 
linn. Bordeaux 20: 561 (1860). Type species: D. 
indicum (L.) Des Moul. - Pyrethrum sect. 
Dendranthema DC. 

Perennial herbs or half-shrubs. Leaves alternate, pinnatisect, 
lobed, serrate or rarely entire. Capitula laxly corymbose or 
solitary, radiate. Involucral bracts generally with dark brown 
margins. Receptacle convex to conical, epaleate. Ray florets 
female, fertile; limb white, pink, or yellowish. Disc corolla 
5-lobed; tube generally with sessile glands. Apical anther 
appendage subtriangular. Cypselas obovoid, faintly 5-8- 
ribbed, thin-walled, generally with myxogenic cells in rows. 
Pappus absent. 

Distribution. Asia, mainly in China and Japan, one species 
extending to E. Europe (D. zawadskii). - 37 spp. 

Dendranthema differs from Tanacetum (i.e. Chrysanthemum 



114 



K. BREMER AND C. J. HUMPHRIES 



s. 1.) by its obovoid, thin-walled, generally myxogenic cypse- 
las without a pappus. It was revised by Tzvelev (in Komarov, 
1961). Later Tzvelev (1985) transferred three species of 
Dendranthema to a new genus Arctanthemum and later 
erected the new genus Hulteniella based on Arctanthemum 
integrifolium (Richardson) Tzvelev (Tzvelev, 1987). Other 
related genera are Brachanthemum and Tridactylina. 

There is still much work to be undertaken on this species- 
rich and horticulturally interesting genus. There are many, 
poorly understood species, known only from the original 
description, and probably several undescribed ones. Possibly 
there are other species to be transferred from Tanacetum. 
Future revision should be undertaken with Ajania, since 
some species of Dendranthema may be more closely related 
to Ajania than to other species within Dendranthema. In 
other words, Dendranthema may be paraphyletic when Aja- 
nia and its relatives (i.e. Artemisia etc.) are excluded. As 
presently understood, Dendranthema is always radiate and 
Ajania always disciform. Most species of Dendranthema also 
have involucral bracts with dark brown margins, a character 
that may have been secondarily lost in the small-headed 
species of Ajania. 

The well-known autumn-flowering chrysanthemums of 
horticulture are derived from D. grandiflorum and D. indi- 
cum. 

The list of species is compiled mainly from Flora URSS 
(Komarov, 1961) and the Flora of the People's Republic of 
China (Ling Sl Shih, 1980, 1983; Shih & Fu, 1983), as well as 
from accounts from Japan by Kitamura (1940, 1978, 1979). 
These authors transferred a number of species from Chrysan- 
themum to Dendranthema. Kitamura also included four disci- 
form species, D. pallasianum, D. rupestre, D. pacificum, and 
D. shiwogiku, in section Ajania. The latter we consider as a 
separate genus in agreement with various other authors and 
so the disciform species are here listed under Ajania. The 
generic distinction, as discussed under that genus, is in doubt 
but pending a detailed study of both genera we consider it 
best to transfer a few disciform Japanese Dendranthema 
species to Ajania rather than to recombine the numerous 
Chinese Ajania species under Dendranthema. 

*D. aphrodite (Kitam.) Kitam. Japan. 

*D. argyrophyllum (Ling) Ling «fe Shih. China. 

D. arisanense (Hayata) Ling & Shih. Taiwan. 

D. boreale (Makino) Ling ex Kitam. China, Korea, Japan. 

D. chalchingolicum (Grubov) Bremer & Humphries, comb. 

nov. Basionym: Chrysanthemum chalchingolicum Grubov 

in Bot. Zhurn. 15: 1592 (1972). Mongolia. 
D. chanetii (A. Leveille) Shih {D. erubescens (Stapf) 

Tzvelev). China, Korea. 
D. coreanum (A. Leveille & Vaniot) Vorosch. Korea. 
*D. crassum (Kitam.) Kitam. Japan. 
D. cuneifolium (Kitam.) Bremer & Humphries, comb. nov. 

Basionym: Chrysanthemum cuneifolium Kitam. in Acta 

phytotax. geobot. Kyoto 7: 68 (1938). Japan. 
*D. dichrum Shih. China. 
*D. glabriusculum (W. Smith) Shih. China. 
D. grandiflorum (Ramat.) Kitam. (D. morifolium (Ramat.) 

Tzvelev). China, much cultivated. 
D. hypargyrum (Diels) Ling & Shih. China. 
D. indicum (L.) Des Moul. China, Korea, Japan, much 

cultivated. 
D. japonense (Nakai) Kitam. Japan. 
D. japonicum (Makino) Kitam. Japan. 



D. lavandulifolium (Fischer ex Trautv.) Kitam. China. 

*D. littorale (Mackawa) Tzvelev. Far East, Japan. 

D. maximowiczii (V. Komarov) Tzvelev. Far East, China, 
Korea. 

D. miyatojimense (Kitam.) Hind. Japan. 

D. mongolicum (Ling) Tzvelev. E. Siberia, China. 

*D. morii (Hayata) Kitam. Taiwan. 

D. nankingense (Hand.-Mazz.) Y. R. Ling. China. 

*D. okiense (Kitam.) Kitam. Japan. 

D. oreastrum (Hance) Ling (D. sichotense Tzvelev). Far East, 
China, Korea. 

D. ornatum (Hemsley) Kitam. Japan. 

*D. parvifolium (Chang) Shih. China. 

D. potentilloides (Hand.-Mazz.) Shih. China. 

*D. rhombifolium Ling & Shih. China. 

*D. sinchangense (Ueki) Kitam. Korea. 

D. sinuatum (Ledeb.) Tzvelev. C. Asia, MongoHa. 

*D. vestitum (Hemsley) Ling. China. 

*D. weyrichii (Maxim.) Tzvelev. Far East, Japan. 

*D. xeromorphum Khokr. Russia. 

D. yezoense (T. Maek.) Hind. Japan. 

D. yoshinaganthum (Makino ex Kitam.) Kitam. Japan. 

D. zawadskii (Herbich) Tzvelev (D. naktongense (Nakai) 
Tzvelev). From E. Europe through Russia to China, Mon- 
golia, and Japan. 



32. ARCTANTHEMUM (Tzvelev) Tzvelev in Nov. 
Sist. Vysshikh Rast. 22: 274 (1985). Type species: A. 
arcticum (L.) Tzvelev (including Hulteniella 
Tzvelev). 

Perennial rhizomatous herbs. Leaves rosulate to alternate, 
obovate-spathulate to linear, apically lobed-serrate or entire. 
Capitula solitary, radiate. Involucral bracts with dark brown 
margins. Receptacle convex to conical, epaleate. Ray florets 
female, fertile; limb white. Disc corolla 5-lobed; tube gener- 
ally with sessile glands. Apical anther appendage subtriangu- 
lar. Cypselas oblong, somewhat 5-8-ribbed, without 
myxogenic cells. Pappus absent. 

Distribution. Arctic Eurasia, Siberia, Far East, Japan, 
Arctic N. America. - 4 spp. 

Arctanthemum was formerly a section of Dendranthema. It 
consists of more or less rosulate herbs with an arctic distribu- 
tion, whereas most Dendranthema species are leafy herbs or 
herbaceous perennials from China and Japan. It is worth 
noting that Tzvelev (1987) transferred A. integrifolium to a 
new genus, Hulteniella Tzvelev, which we do not recognize 
here. 

A. arcticum (L.) Tzvelev {Dendranthema arcticum (L.) 
Tzvelev). Far East and Arctic America. 

A. hultenii (A. & D. Love) Tzvelev {Dendranthema hultenii 
(A. & D. Love) Tzvelev). Arctic Eurasia and Arctic 
America. 

A. integrifolium (Richardson) Tzvelev {Dendranthema inte- 
grifolium (Richardson) Tzvelev, Hulteniella integrifolium 
(Richardson) Tzvelev). NE Siberia and Arctic N. America. 

A. kurilense (Tzvelev) Tzvelev {Dendranthema kurilense 
(Tzvelev) Tzvelev). Far East, Japan. 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 

33. TRIDACTYLINA (DC.) Schultz-Bip. in Webb & 
Berthelot, Hist. nat. Iks Canaries 3 (2,2): 245 
(1844). Type species: T. kirilowii (Turcz. ex DC.) 
Schultz-Bip. 

An annual herb. Leaves alternate, few-lobed. Capitula laxly 
corymbose, radiate. Involucral bracts with dark brown mar- 
gins. Receptacle flat to convex, epaleate. Ray florets neuter; 
Hmb yellow. Disc corolla 5-lobed. Apical anther appendage 
subtriangular. Cypselas 5-ribbed, with myxogenic cells and 
with an apical rim. Pappus absent. 

Distribution. E. Siberia. - Monotypic. 

According to Tzvelev in Flora URSS (Komarov, 1961) this 
species is clearly related to Dendranthema but differs by its 
annual habit and neutral rays. It is similar for example in 
foliage to some species of Dendranthema and Arctanthemum. 

34. AJANIA Polj. in Bot. Mater. Gerb. bot. Inst. V. 
A. Komarova 17: 419 (1955). Type species: A. 
pallasiana (Fischer ex Besser) Polj . 

Perennial herbs or half-shrubs. Leaves alternate, pinnatisect, 
lobed, serrate, or rarely entire. Capitula small, corymbose or 
rarely solitary, disciform. Receptacle convex to conical, 
epaleate. Outer female florets in one row. Corollas of central 
florets 5-lobed; tube generally with sessile glands. Apical 
anther appendage subtriangular. Cypselas obovoid, faintly 
4-6-ribbed, thin-walled, generally with myxogenic cells in 
rows. Pappus absent. 

Distribution. C. Asia, mainly in China, also in Japan. - 34 
spp. 

Poljakov removed a number of species from Artemisia to a 
separate genus Ajania. He noted that Ajania has fertile 
florets and the common form of distinct 5-lobed corollas with 
spreading lobes, whereas in Artemisia only some of the florets 
in each capitulum are fertile, producing mature fruits, and the 
corollas have small erect lobes. Ajania also has a corymbose 
inflorescence as opposed to the elongated inflorescences of 
Artemisia. Poljakov speculated ihdii Ajania is a very old group 
derived from the same ancestors as Artemisia. 

Tzvelev (in Komarov, 1961) accepted Ajania although he 
retained some of Poljakov's Ajania species in Artemisia. He 
also noted the close affinity between Dendranthema and 
Ajania, and speculated that Ajania and Artemisia are two 
convergent and habitually similar lines independently 
evolved from the same 'dendranthemoid' ancestors. More 
parsimoniously, Ajania may be considered the plesiomorphic 
sister group to Artemisia and all its relatives. Possibly Ajania 
or part of the genus is the sister group of Artemisia and the 
other genera with smooth or short-spined pollen, as indicated 
in the cladogram. Dendranthema is even more plesiomorphic 
and possibly paraphyletic with Ajania, and Artemisia etc. 
excluded. Ajania differs from Dendranthema by its smaller, 
disciform, densely corymbose capitula. No doubt this species- 
rich and little-known genus deserves a detailed study together 
with Dendranthema. 

A few species of Ajania have recently been placed in a 
separate genus, Phaeostigma. There may be several species of 
Ajania (Muldashev, 1982, 1983) that should be transferred to 
Phaeostigma, should the latter be kept distinct. The type 
species of Ajania, A. pallasiana, and A. latifolia, A. rupestris, 
and A. shiwogiku are all examples of species similar to 



115 

Phaeostigma. The matter is discussed further under Phaeo- 
stigma. 

Tzvelev suspected that the distinction between Ajania and 
Dendranthema would become difficult after examination of 
the extensive Chinese material. Nevertheless Ling & Shih 
(1980, 1983) kept them separate in their accounts of the 
Chinese species. Our list of species is compiled mainly from 
their treatments and from Flora URSS (Komarov, 1961). The 
species from Tibet were listed by Shih & Fu (1979) and the 
Japanese species are from Kitamuras's (1978) Dendranthema 
sect. Ajania (see discussion under Dendranthema). 

A. achilleoides (Turcz.) Polj. ex Grubov. Mongolia. 

A. adenantha (Diels) Ling & Shih. China. 

*A. brachyantha Shih. China. 

A. breviloba (Franchet ex Hand.-Mazz.) Ling &. Shih. China. 

*A. elegantula (W. Smith) Shih. China. 

A. fastigiata (Winkler) Polj. C. Asia, China. 

A. fruticulosa (Ledeb.) Polj. (A. aureoglobosa (W. Smith & 

Farrer) Muld.). C. Asia, E. Siberia, Mongolia, China. 
A. gracilis (Hook. f. & Thomson) Polj. ex Tzvelev. C. Asia, 

China, Himalaya. 
*A. grubovii Muld. MongoHa. 
*A. junnanica Polj. China. 
A. khartensis (Dunn) Shih {A. mutellina (Hand.-Mazz.) 

Muld.). China. 
*A. kokanica (H. Kraschen.) Tzvelev. C. Asia. 
A. latifolia Shih. China. 
A. myriantha (Franchet) Ling ex Shih {A. oresbia (W. Smith) 

Muld.). China. 
M. nana (H. Kraschen.) Muld. China. 
*A. nematoloba (Hand.-Mazz.) Ling & Shih. China. 
*A. nitida Shih. China. 
A. nubigena (Wallich) Shih. China. 
A. paciflea (Nakai) Bremer & Humphries, comb. nov. 

Basionym: Chrysanthemum pacificum Nakai in Bot. Mag., 

Tokyo 42: 462 (1928) {Dendranthema pacificum (Nakai) 

Kitam.). Japan. 
A. pallasiana (Fischer ex Besser) Polj. (Dendranthema palla- 

sianum (Fischer ex Besser) Vorosch.). Far East, China, 

Korea, Japan. 
A. parviflora (Griin.) Ling. China. 
A. potaninii (H. Kraschen.) Polj. China. 
A. przewalskii Polj. China. 
*A. purpurea Shih. China. 

*A. remotipinna (Hand.-Mazz.) Ling & Shih. China. 
*A. roborowskii Muld. China. 
A. rupestris (Matsum. ex Koidz.) Muld. Japan. 
A. scharnhorstii (Regel & Schmalh.) Tzvelev. C. Asia. 
*A. sericea Shih. China. 
A. shiwogiku (Kitam.) Bremer & Humphries, comb. nov. 

Basionym: Chrysanthemum shiwogiku Kitam. in Acta phy- 

totax. geobot. Kyoto 4: 71 (1935). Japan. 
A. tenuifolia (Jacquem.) Tzvelev. Himalaya. 
A. tibetica (Hook. f. & Thomson) Tzvelev. C. Asia, China, 

Himalaya. 
*A. trilobata Polj. C. Asia. 
*A. tripinnatisecta Ling & Shih. China. 

35. PHAEOSTIGMA Muld. in Bot. Zhurn. 66: 586 
(1981). Type species: P. salicifolium (Mattf.) Muld. 

Perennial herbs or half-shrubs. Leaves alternate, pinnatifid to 
shallowly lobed or entire. Capitula small, densely corymbose, 



116 



K. BREMER AND C. J. HUMPHRIES 



disciform. Receptacle convex, epaleate. Outer female florets 
in one row. Central floret corolla 5-lobed; lobes erect. Apical 
anther appendage subtriangular. Style-branches brownish. 
Cypselas 4-6-ribbed; ribs projected into minute teeth. Pap- 
pus absent. 

Distribution. China. - 3 spp. 

Muldashev made a detailed comparison of Ajania fruticulosa 
and the three species of Ajania transferred by him to Phaeo- 
stigma. According to Muldashev Phaeostigma is distinguished 
by its brownish style-branches and erect corolla lobes. The 
latter character, as well as the suffruticose habit and the less 
spiny pollen, are characters similar to those of Artemisia. 
Muldashev stated that phylogenetically Phaeostigma is 
related to the ancestors of Ajania and distantly related to 
Artemisia, mainly on account of its pollen morphology. 

The brownish style-branches is a difficult character to 
evaluate. Similar style-branches may be present in related 
genera. Although Shih & Fu (1983) reduced Phaeostigma to 
synonomy under Ajania we have kept them distinct. In that 
case Phaeostigma may be another grade group between 
Ajania and Artemisia and its relatives. The actual circum- 
scription of Phaeostigma will have to be assessed in a wider 
study of Ajania and especially those Ajania species that are 
similar in habit to Phaeostigma. 

P. quercifolium (W. Smith) Muld. (Ajania quercifolia (W. 

Smith) Ling & Shih) 
P. saUcifolium (Mattf.) Muld. (Ajania salicifoUa (Mattf.) 

Polj.) 
P. variifolium (Chang) Muld. (Ajania manshurica Polj., 

Ajania ramosa (Chang) Shih, Ajania variifolia (Chang) 

Tzvelev) 

36. STILPNOLEPIS H. Kraschen. in Nov. Sist. 
Vysshikh Rast. 2: 207 (1946). Type species: S. 
centiflora (Maxim.) H. Kraschen. - Elachanthemum 
Ling & Y. R. Ling. 

Annual or perennial herbs. Leaves opposite or alternate, 
pinnatisect, few-lobed or entire. Capitula laxly corymbose, 
discoid. Involucral bracts widely obovate and largely scari- 
ous. Receptacle convex-subconical, epaleate. Corolla 
5-lobed, with a distinct tube and more or less crateriform 
limb, glandular. Apical anther appendage acuminate- 
triangular. Cypselas obHquely obovoid to narrowly obovoid- 
oblanceolate, thin-walled, without ribs, densely glandular. 
Pappus absent. 

Distribution. Mongolia and China. - 2 spp. 

According to Krascheninnikov (1946) S. centiflora resembles 
Artemisia, but does not share the obovoid cypselas, the 
slender corollas, the acuminate anther appendages and the 
paniculate inflorescence of Artemisia and closely related 
genera. Though elongated, the cypselas are basically the 
same as in other genera of subtribe Artemisiinae; obovoid 
(oblanceolate), thin-walled and devoid of ribs. There is no 
doubt about the subtribal position of Stilpnolepis . 

According to Ling & Y. R. Ling (1978) the second species, 
5. intricata, formerly classified in Artemisia or Seriphidium, 
should be separated from those genera because of the long 
peduncles and the laxly corymbose inflorescence, the cen- 
trally abortive disc florets, and the multicostate appearance 
of the rows of myxogenic cells on the cypselas. Hence they 



created a new genus Elachanthemum for this species. How- 
ever, Ling and Y. R. Ling did not consider a possible 
relationship with Stilpnolepis. Recently Shih (1985) argued 
that Elachanthemum intricatum is closely related to Stilpnol- 
epis centiflora and his treatment is followed here. Both 
species have discoid and laxly corymbose capitula, most 
parsimoniously interpreted as apomorphic within the Artemi- 
sia group, and hence synapomorphies uniting the two species. 

S. centiflora (Maxim.) H. Kraschen. 

S. intricata (Franchet) Shih (Elachanthemum intricatum 
(Franchet) Ling & Y. R. Ling 

37. AJANIOPSIS Shih in Acta phytotax. sin. 16: 86 
(1978). Type species: A. penicilliformis Shih. 

A pubescent annual herb. Leaves alternate, pinnatisect and 
few-lobed. Capitula rather small, few together in dense 
corymbs, disciform. Receptacle convex, epaleate. Outer 
female florets in one row, tapering above, apically densely 
pilose with erect, straight hairs. Central floret corolla 
5-lobed, apically densely pilose with erect, straight hairs. 
Apical anther appendage subtriangular. Cypselas obovoid, 
thin-walled with 3-6 rows of myxogenic cells. Pappus absent. 

Distribution. China, Tibet. - Monotypic. 

This is a distinctive species and undoubtedly a member of 
Artemisiinae, though its immediate relatives are difficult to 
indicate. Shih (1978) compared it to both Ajania and Artemi- 
sia. It is distinguished by its annual habit, the densely 
corymbose capitula and apically pilose corollas. The genus 
differs from Ajania only by the corollas being pilose at the 
apex. The cypsela ribs, described and illustrated by Shih, 
appear to be rows of myxogenic cells. We have only exam- 
ined the holotype and avoided consuming type material for a 
detailed cypsela investigation. 

38. FILIFOLIUM Kitam. in Acta phytotax. geobot. 
Kyoto 9: 157 (1940). Type species: F. sibiricum (L.) 
Kitam. 

A perennial herb, basally somewhat woody and covered with 
fibrous leaf sheaths. Leaves alternate, pinnatisect with long 
fihform lobes. Capitula rather small, corymbose, disciform. 
Receptacle conical, epaleate. Outer female florets tapering 
above, minutely 4-dentate. Central florets apparently her- 
maphrodite but functionally male and female-sterile; corollas 
compressed together in a resinous mass, 5-lobed. Apical 
anther appendage triangular. Cypselas obliquely obovoid, 
thin-walled, with 2 adaxial-lateral rows of myxogenic cells. 
Pappus absent. 

Distribution. Far East, China and Korea. - Monotypic. 

Filifolium was earlier included in Artemisia sect. Dracunculus 
on account of the female-sterile disc florets. However, the 
inflorescence is corymbose, the style is distinctly divided and 
the apical appendages of the anthers are triangular, not 
linear-lanceolate. In habit Filifolium is more similar to some 
species of Ajania than to Artemisia. In pollen and cypsela 
characters Filifolium is apomorphic compared to Ajania. 
Hence, it occupies an intermediate position within the sub- 
tribe, as expressed in the cladogram. The obliquely obovoid 
cypselas with two distinct rows of myxogenic cells appear to 
be autapomorphic for this genus. 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



117 



39. SPHAEROMERIA Nutt. in Trans. Am. phil. Soc. 
2 (7): 401 (1841). Type species: S. capitata Nutt. - 
Chamartemisia Rydb. - Vesicarpa Rydb. 

Perennial herbs or half-shrubs, sometimes rather compact 
and basally woody. Leaves alternate to rosulate, pinnatisect 
and few-lobed or entire. Capitula apparently corymbose but 
sometimes somewhat paniculate or capitate or soHtary, disci- 
form. Receptacle flat to conical, epaleate, rarely pubescent. 
Outer female florets tapering above, sometimes glandular, 
rarely with dolabriform hairs at the apex. Central florets with 
corolla 5-lobed, sometimes glandular, rarely with dolabri- 
form hairs at the apex. Apical anther appendage subtriangu- 
lar. Cypselas obovoid-oblong, sometimes faintly ribbed, 
sometimes with myxogenic cells, often with an apical rim. 
Pappus usually absent or rarely minutely coroniform, with 
small scales. 

Distribution. Western N. America in California, Colo- 
rado, Montana, Nevada, Oregon, Utah and Wyoming, and in 
Mexico. - 9 spp. 

Sphaeromeria was originally proposed by Nuttall to circum- 
scribe 5. capitata and 5. argentea. Torrey & Gray (1843) 
reduced Sphaeromeria to a section of Tanacetum simply on 
combination of three plesiomorphic characters, the yellow 
5-lobed corollas, the woody base and the alternate leaves. 
Rydberg (1916) returned Sphaeromeria to its generic status 
and erected two new genera, Vesicarpa to include Artemisia 
potentilloides A. Gray with a hairy receptacle and Chamarte- 
misia for Tanacetum compactum Hall with pappose cypselas. 

A. Holmgren et al. (1976) presented a detailed study of the 
three genera Sphaeromeria, Vesicarpa and Chamartemisia, 
and concluded that they are more closely related to one 
another than to any other group. Sphaeromeria appears to be 
more closely related to Artemisia than Tanacetum. However, 
of the 12 characters discussed by A. Holmgren et al. not one 
is autapomorphic for Sphaeromeria. They are present in all 
other genera of Artemisiinae. The presence of a small 
coroniform pappus in one species is an unusual feature within 
Artemisiinae but a parallel case is Crossostephium. 

Sphaeromeria is similar in habit to Kaschgaria. The inflo- 
rescences of Sphaeromeria are variable and the transforma- 
tions within the genus are difficult to assess. In the cladogram 
the corymbose condition is considered plesiomorphic but 
there are species with slightly elongated inflorescences resem- 
bling those of Kaschgaria. The two genera are possibly 
closely related. 

The list of species is taken from A. Holmgren et al., who 
also provided a key. 

S. argentea Nutt. 

S. cana (D. C. Eaton) A. A. Heller 

5. capitata Nutt. 

S. compacta (H. M. Hall) A. Holmgren, Shultz & Lowrey 
{Chamartemisia compacta (H. M. Hall) Rydb.) 

S. diversifolia (D. C. Eaton) Rydb. 

*S. martirensis (Wiggins) A. Holmgren, Shultz & Lowrey 

S. potentilloides (A. Gray) A. A. Heller {Vesicarpa potentil- 
loides (A. Gray) Rydb.) 

5. ruthiae A. Holmgren, Shultz & Lowrey 

5. simplex (Nelson) A. A. Heller 



40. KASCHGARIA Polj. in Bot. Mater. Gerb. bot. 
Inst. V. A. Komarova 18: 282 (1957). Type species: 
K. brachanthemoides (Winkler) Polj. 

Rather woody half-shrubs; indumentum of stellate hairs. 
Leaves alternate, entire or few-lobed. Capitula rather small 
and few in an elongated panicle, at the summit fasciculate, 
disciform. Receptacle conical, epaleate. Outer female florets 
few, tapering above. Central floret corolla apically with 
stellate hairs, 5-lobed. Apical anther appendage linear- 
lanceolate. Cypselas obovoid, thin-walled. Pappus absent. 

Distribution. C. Asia and China. - 2 spp. 

Poljakov removed the two species of Kaschgaria from Arte- 
misia mainly because of their stellate corolla hairs. He also 
stated that Kaschgaria is closer to Seriphidium than to 
Artemisia s. s., but Kaschgaria differs from Seriphidium, for 
example, by its disciform (heterogamous) versus a discoid 
(homogamous) capitulum. The possible relationship to 
Sphaeromeria (then part of Tanacetum) was not mentioned 
by Poljakov, who probably did not consider the North 
American species of Tanacetum. He closed his discussion 
noting that the systematic position of Kaschgaria could not be 
settled before the whole Asian part of the Artemisia group 
had been properly considered. We agree that Kaschgaria is a 
distinct genus probably related to Seriphidium, or possibly 
more closely to Sphaeromeria, although there is no obvious 
synapomorphy uniting the two genera. 

K. brachanthemoides (Winkler) Polj . 

K. komarovii (H. Kraschen & N. Rubtzov) Polj. 

41. SERIPHIDIUM (Besser ex Hook.) Fourr. in 
Annls. Soc. linn. Lyon. II, 17: 89 (1869) Type 
species: S. maritimum (L.) Polj. -Artemisia sect. 
Seriphidium Besser - Artemisiastrum Rydb. 

Perennial herbs, half-shrubs or annual herbs. Leaves alter- 
nate, pinnatisect. Capitula small and few-flowered, oblong, 
numerous in a long panicle, discoid. Involucral bracts in 4-7 
rows, unequal; the outer short and rounded, the inner 
gradually longer and linear. Receptacle small, conical, epale- 
ate or occasionally paleate. Corolla 5-lobed, tubular, yellow 
to purple. Apical anther appendage linear-lanceolate. Cypse- 
las small, obovoid to ellipsoid, somewhat flattened or trique- 
trous, thin-walled. Pappus absent. 

Distribution. From Europe throughout temperate Asia to 
N. America (5 spp.), though mainly in central Asia. - 134 
spp. 

Seriphidium, established as a genus by Poljakov (1961), is 
almost always considered a section or subgenus of Artemisia 
(e.g. Ward, 1940). We agree with Poljakov (1961), Leonova 
(1970) and Filatova (1981, 1982fl, b 1984) that it should be 
kept distinct. Seriphidium differs in a number of characters, 
representing synapomorphies. Furthermore, its sister group 
appears to be found outside Artemisia s. s., and Seriphidium 
cannot be considered a specialized segregate of Artemisia. 
Seriphidium is most closely related either to a larger group of 
genera including Artemisia or some other genus (or genera), 
e.g. Kaschgaria, within the Artemisiinae. The genus is char- 
acterized by its discoid, homogamous capitula, not disciform 
and heterogamous as in Artemisia. The most parsimonious 
interpretation for this character is to assume that the outer 



118 



K. BREMER AND C. J. HUMPHRIES 



female florets present in related genera have been lost. This 
was originally suggested by Krascheninnikov (1946)'although 
hotly contested by Poljakov (1961). It must be noted that this 
character breaks down in 5. bigelowii. This species is usually 
included in Artemisia section Abrotanum because there are at 
least some capitula within inflorescences that have one or two 
female outer florets. However, on the basis of involucral 
(many-rowed bracts) and anther (slender apical appendages) 
characters S. bigelowii is certainly a member of Seriphidium. 
It is also virtually identical to the American 5. tridentatum, 
which is homogamous and always included in Seriphidium 
(see Weber, 1984). Heterogamy in 5. bigelowii is plesiomor- 
phic or a secondary reversal from the homogamous condi- 
tion, which then can be interpreted as a synapomorphy for 
the genus. 

The corollas of Seriphidium are tubular and the teeth 
usually infolded for much of anthesis. During anthesis the 
short flat lobes of the style are often still together and 
enclosed in the anther tube, a feature mostly associated with 
cleistogamy. Mature cypselas are hard to find and there are 
many occasions when only one will mature in a head of 3-10 
florets. 

The apical anther appendages of Seriphidium are very 
slender, narrowly lanceolate to linear and notably different 
from those of Artemisia and related genera. The involucre is 
specialized with 4-7 rows of overlapping bracts, as compared 
to the 2^-rowed involucre of related genera. 

The monotypic Artemisiastrum {A. palmeri=S. palmeri) 
was separated by Rydberg (1916) on the presence of recep- 
tacular scales. Parallel cases are legion in the Anthemideae 
and in Asteraceae as a whole. Here the presence of paleae is 
best considered a gain character, a feature also noted by Hall 
& Clements (1923). In all characters S. palmeri is virtually 
identical to other taxa within Seriphidium. 

The list of species has been compiled from the standard 
floras, cf. under Artemisia, and by original research with Y. 
R. Ling (1991fl, b) during his sabbatical year at The Natural 
History Museum, London. Except for cases of basionyms, 
synonymous Artemisia names are not included when the same 
epithet is involved. Also, combinations from Artemisia by 
Bremer & Humphries have been validly pubhshed in Ling 
(1991fl). 

5. algeriense (Filat.) Y. R. Ling. Algeria. 

*S. amoenum (Polj.) Polj. C. Asia. 

5. aralense (H. Kraschen.) Polj. SW C. Asia. 

5. arbusculum (Nutt.) W. A. Weber. W. United States. 

5. arenicolum (H. Kraschen. ex Polj.) Y. R. Ling. Afghani- 
stan, Iran, C. Asia. 

*S. argilosum (Beetle) Bremer & Humphries, comb. nov. 
Basionym: Artemisia argilosa Beetle, Rhodora 61: 84 
(1959). United States in Colorado. 

*5. assurgens (Filat.) Bremer & Humphries in Y. R. Ling. 
Russia. 

*S. aucheri (Boiss.) Ling & Y. R. Ling. Iran, Afghanistan, 
China. 

S. badhysi (Krasch. & Lincz. ex Polj.) Polj. C. Asia. 

5. balchanorum (H. Kraschen.) Polj. SW C. Asia. 

*S. baldshuanicum (H. Kraschen. & Zaprj.) Polj. Afghani- 
stan, C. Asia. 

S. barrelieri (Besser) Sojak. Spain, N. Africa. 

S. bicolor (Rech. f. & Wagenitz) Bremer & Humphries, 
comb. nov. Basionym: Artemisia bicolor Rech. f. & 
Wagenitz in Anz. ost. Akad. Wiss. Mathematische Natur- 



wissenschaftliche Klasse., 98: 78 (1961). Afghanistan. 
S. bigelowii (A. Gray) Bremer & Humphries, comb. nov. 

Basionym: Artemisia bigelowii A. Gray in Torrey Pacific R. 

Rep. 4: 110 (1857). S. and W. United States. 
*S. borotalense (Polj.) Ling & Y. R. Ling. C. Asia, China. 
5. botschantzevii (Filat.) Y. R. Ling. Russia. 
*S. brevifolium (Walhch ex DC.) Ling & Y. R. Ling. 

Afghanistan, Pakistan, India, China. 
S. caerulescens (L.) Sojak. S. Europe. 
*5. camelorum (H. Kraschen.) Polj. C. Asia. 
5. canum (Pursh) W. A. Weber. W. North America. 
S. chitralense (Podl.) Bremer & Humphries, comb. nov. 

Basionym: Artemisia chitralensis Podl., Fl. iranica 158: 198 

(1986). Afghanistan, Pakistan. 
*5. ciniforme (H. Kraschen. & Popov ex Polj.) Polj. Iran, C. 

Asia. 
S. cinum (P. Bergius ex Polj.) Polj. C. Asia, China. 
S. compactum (Fischer ex DC.) Polj. C. Asia, S. Siberia, 

China in Sinkiang, MongoUa. 
*S. cretaceum (Fiori) Bremer & Humphries, comb. nov. 

Basionym: Artemisia caerulescens var. cretacea Fiori in 

Fiori & Paoletti, Fl. Italia 3: 251 (1904). S. Europe in Italy. 
5. densifolium (Filat.) Y. R. Ling. Algeria 
5. deserti (H. Kraschen.) Polj. Afghanistan, Iran, SW C. 

Asia. 
5. diffusum (H. Kraschen. ex Polj.) Y. R. Ling. Iran, 

Afghanistan, Pakistan. 
*5. dubjanskyanum (H. Kraschen. ex Polj.) Polj. C. Asia. 
S. dumosum (Polj.) Polj. C. Asia. 
*5. dzevanovskyi (Leonova) Sojak. Krym. 
*5. elongatum (Filat. & Ladyg.) Bremer & Humphries in Y. 

R. Ling. C. Asia. 
*5. eremophilum (H. Kraschen. & Butkov ex Polj.) Bremer 

& Humphries in Y. R. Ling. C. Asia. 
5. federovii (Rzazade) Y. R. Ling. C. Asia. 
*S. fedtschenkoanum (H. Kraschen.) Polj. C. Asia, China in 

Xinjiang. 
*S. ferganense (H. Kraschen. ex Polj.) Polj. C. Asia. 
*5. finitum (Kitagawa) Ling & Y. R. Ling. China. 
5. fragrans (Willd.) Polj. Afghanistan, Caucasus, Iran, C. 

Asia. 
S. freitagii (Podl.) Bremer & Humphries, comb. nov. 

Basionym: Artemisia freitagii Podl., Fl. iranica 158: 193 

(1986). Afghanistan. 
*5. fulvellum (Filat. & Ladyg.) Bremer & Humphries in Y. 

R. Ling. C. Asia. 
S. ghazniense (Podl.) Bremer & Humphries, comb. nov. 

Basionym: Artemisia ghazniensis Podl., Fl. iranica 158: 213 

(1986). Afghanistan. 
S. ghoratense (Podl.) Bremer & Humphries, comb. nov. 

Basionym: Artemisia ghoratensis Podl., Fl. iranica 158: 

197. Afghanistan. 
*S. glanduligerum (H. Kraschen. ex Polj.) Polj. Afghanistan, 

Pakistan, C. Asia. 
*5. glaucinum (H. Kraschen. ex Polj.) Bremer & Humphries 

in Y. R. Ling. C. Asia. 
5. gorjaevii (Polj.) Y. R. Ling. C. Asia. 
S. gracilescens (H. Kraschen. & Iljin) Polj. C. Asia, S. 

Siberia, Mongolia, China. 
S. grenardii (Franchet) Y. R. Ling & Humphries. China. 
*S. gurganicum (H. Kraschen.) Bremer & Humphries in Y. 

R. Ling. C. Asia. 
*S. gypsaceum (H. Kraschen., Popov & Lincz. ex Polj.) Polj. 

Iran, SW & C. Asia. 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



119 



*S. halophilum (H. Kraschen.) Polj. C. Asia. 

*S. heptapotamicum (Polj.) Ling & Y. R. Ling. C. Asia, 

China. 
S. herba-album (Asso) Sojak. SW Europe, Turkey, Middle 

East, Iran, Himalayas. 
S. incultum (Del.) Y. R. Ling. Egypt. 
S. issykkulense (Polj.) Polj. C. Asia, China. 
S. junceum (Karelin & Kir.) Polj. C. Asia, China in Xinjiang. 
S. kandaharense (Podl.) Bremer & Humphries, comb. nov. 

Basionym: Artemisia kandaharensis Podl., Fl. iranica 158: 

217. Afghanistan. 
*S. karatavicum (H. Kraschen. & Abolin ex Polj.) Ling & Y. 

R. Ling. C. Asia, China. 
*S. kasakorum (H. Kraschen.) Bremer & Humphries, comb. 

nov. Basionym: Artemisia maritima ssp. kasakorum H. 

Kraschen., Otch. Rab. Pochv.-Bot. Otr. Kazakhst. Exped. 

Akad. Nauk SSSR 4(2): 272 (1930). C. Asia. 
*S. kaschgaricum (H. Kraschen.) Polj. C. Asia, China in 

Xinjiang. 
*S. kemrudicum (H. Kraschen.) Polj. SW & C. Asia. 
S. kermanense (Podl.) Bremer & Humphries, comb. nov. 

Basionym: Artemisia kermanensis Podl., Fl. iranica 158: 

206 (1986). Iran. 
*S. khorassanicum (Podl.) Bremer & Humphries, comb. nov. 

Basionym: Artemisia khorassanica Podl., Fl. iranica 158: 

210 (1986). Iran, Afghanistan. 
*S. knorringianum (H. Kraschen.) Polj. C. Asia. 
*S. kochiiforme (H. Kraschen. & Lincz. ex Polj.) Polj. 

Afghanistan, C. Asia. 
*S. kopetdaghense (H. Kraschen. ex Polj.) Polj. Afghanistan, 

Iran, Afghanistan, SW C. Asia. 
S. korovinii (Polj.) Polj. Afghanistan, C. Asia. 
5. korshinskyi (H. Kraschen. ex Polj.) Y. R. Ling. Afghani- 
stan. 
S. kurramense (Qaz.) Y. R. Ling. Afghanistan, Pakistan. 
*S. lehmannianum (Bunge) Polj. Afghanistan, C. Asia. 
S. lerchianum (G. Weber in Stechm.) Polj. From SE Europe 

in Bulgaria through Russia and C. Asia to S. Siberia. 
S. lessingianum (Besser) Polj. SE Russia, C. Asia, S. Siberia. 
S. leucodes (Schrenk) Polj. C. Asia. 
S. leucotrichum (H. Kraschen. ex Polj.) Bremer & 

Humphries in Y. R. Ling. Afghanistan, Pakistan, C. Asia. 
*S. longilobum (Osterh.) Bremer & Humphries, comb. nov. 

Basionym: Artemisia spiciformis var. longiloba Osterh. in 

Muhlenbergia 4: 69 (1908). N. America. 
S. maritimum (L.) Polj. W., N. and E. Europe, Iran, C. Asia 

and the Himalayas. 
S. mendozanum (DC.) Bremer & Humphries, comb. nov. 

Basionym: Artemisia mendozana DC, Prodr. 6: 105 

(1837). W. North America. 
*S. minchunense Ling & Y. R. Ling. China. 
S. mogoltavicum (Polj.) Y. R. Ling. C. Asia. 
*S. mongolorum (H. Kraschen.) Ling & Y. R. Ling. China in 

Mongolia. 
S. mucronulatum (Polj.) Y. R. Ling. C. Asia. 
*S. namanganicum (Polj.) Polj. C. Asia. 
*S. nigricans (Filat. & Ladyg.) Bremer & Humphries in Y. R. 

Ling. C. Asia. 
*S. nitrosum (G. Weber ex Stechm.) Polj. SE Russia, C. 

Asia, S. Siberia, China in Xinjiang. 
*S. novum (Nelson) W. A. Weber. W. United States. 
S. nutans (Willd.) Sojak. SE Russia. 
*S. oliverianum (Gay ex Besser) Bremer & Humphries in Y. 

R. Ling. Iran, Afghanistan, Pakistan, C. Asia. 



S. oranense (Deb. ex Filat.) Y. R. Ling. NW Africa (Alge- 
ria). 
5. oratense (Deb. & Filat.) Y. R. Ling. Algeria. 
S. palmeri (A. Gray) Bremer & Humphries, comb. nov. 

Basionym: Artemisia palmeri A. Gray in Proc. Am. Acad. 

11: 79 (1876) (Artemisiastrum palmeri (A. Gray) Rydb.). 

N. America in CaUfornia and Baja California (Mexico). 
5. pauciflorum (G. Weber in Stechm.) Polj. SE Russia, C. 

Asia, S. Siberia. 
S. poljakovii (Filat.) Y. R. Ling. Russia. 
S. polystichum (Polj.) Y. R. Ling. C. Asia. 
S. porrectum (H. Kraschen. ex Polj.) Polj. C. Asia. 
*S. prasinum (H. Kraschen. ex Polj.) Polj. Afghanistan, C. 

Asia. 
*S. prolixum (H. Kraschen. ex Polj.) Polj. C. Asia. 
*S. pygmaeum (A. Gray) W. A. Weber. W. United States. 
*S. quettense (Podl.) Bremer & Humphries, comb. nov. 

Basionym: Artemisia quettensis Podl., Fl. iranica 158: 212 

(1986). Iran, Pakistan. 
S. rhodanthum (Rupr.) Polj. C. Asia. 
*S. rigidum (Nutt.) W. A. Weber. W. United States. 
S. rothrockii (A. Gray) W. A. Weber. W. United States. 
S. saharum (Pomel) Y. R. Ling. Algeria, Tunisia. 
*S. saissanicum (H. Kraschen.) Bremer &. Humphries in Y. 

R. Ling. C. Asia. 
S. santolinum (Schrenk) Polj. (5. lobulifolium (Boiss.) Polj.). 

Iran, C. Asia. 
S. santonicum (L.) Sojak (5. monogynum (Waldst. & Kit.) 

Polj.). SE and E. Europe to C. Asia, Turkey. 
S. sawanense Y. R. Ling & Humphries. China. 
5. schrenkianum (Ledeb.) Polj. C. Asia, S. Siberia, China in 

Xinjiang, Mongolia. 
*5. scopiforme (Ledeb.) Polj. C. Asia, S. Siberia. 
*S. scotinum (Nevski) Polj. Afghanistan, C. Asia. 
*S. semiaridum (H. Kraschen. & Lavrenko) Ling & Y. R. 

Ling. C. Asia. 
S. serotinum (Bunge) Polj. C. Asia. 
*S. sieberi (Besser) Bremer & Humphries in Y. R. Ling. 

Middle East, Iraq, Iran, Afghanistan, SW C. Asia. 
S. skorniakowii (Winkler) Bremer & Humphries in Y. R. 

Ling. C. Asia. 
*5. spicigerum (Koch) Polj. Turkey, Caucasus, Iran. 
*5. stenocephalum (H. Kraschen. ex Polj.) Polj. Afghanistan, 

Pakistan, C. Asia. 
*S. subchrysolepis (Filat.) Bremer & Humphries, comb. nov. 

Basionym: Artemisia subchrysolepis Filat. in Nov. Sist. 

Vysshikh Past. 18: 224 (1981). C. Asia. 
S. sublessingianum (Kell.) Polj. C. Asia, S. Siberia, Mongo- 
lia. 
*S. subsalsum (Filat.) Bremer & Humphries in Y. R. Ling. 

C. Asia. 
5. szowitzianum (Besser) Polj. Caucasus. 
*S. tauricum (Willd.) Polj. Krym, Caucasus, Turkey. 
*S. tecti-mundii (Podl.) Bremer & Humphries, comb. nov. 

Basionym: Artemisia tecti-mundii Podl., Fl. iranica 158: 206 

(1986). Afghanistan, Pakistan. 
S. tenuisectum (Nevski) Polj. C. Asia. 
S. terrae-albae (H. Kraschen.) Polj. C. Asia, Mongolia, 

China. 
*S. thomsonianum (C. B. Clarke) Ling & Y. R. Ling. China. 
S. tianshanicum (H. Kraschen.) Y. R. Ling. C. Asia, China in 

Xinjiang, Mongolia. 
S. transiliense (Polj.) Polj. C. Asia. 
S. tridentatum (Nutt.) W. A. Weber. W. North America. 



120 



K. BREMER AND C. J. HUMPHRIES 



S. tripartitum (Rydb.) W. A. Weber. W. United States. 

*S. turanicum (H. Kraschen.) Polj. Afghanistan, Iran, Paki- 
stan, C. Asia. 

*S. turcomanicum (Gand.) Polj. Iran, SW & C. Asia. 

*S. vachanicum (H. Kraschen. ex Polj.) Polj. Afghanistan, 
Pakistan, C. Asia. 

*S. validum (H. Kraschen. ex Polj.) Polj. C. Asia. 

S. valesianum (Lam.) Y. R. Ling. C. Europe. 

5. vallesiacum (All.) Sojak. S. Europe. 

*S. vaseyanum (Rydb.) W. A. Weber. W. United States. 

42. CROSSOSTEPHIUM Less, in Linnaea 6: 220 
(1831). Type species: C. artemisioides Less. (C. 
chinense (L.) Makino). 

A tomentose shrub. Leaves alternate, narrowly spathulate, 
apically few-lobed or entire. Capitula rather small and 
rounded, paniculate, disciform. Outer involucral bracts 
tomentose, inner scarious. Receptacle hemispherical, epale- 
ate. Outer female florets tubular, 2-3-lobed, glandular. Cen- 
tral florets with corolla tubular, 5-lobed, glandular. Apical 
anther appendage subtriangular. Cypselas obovoid, weakly 
5-ribbed, glandular. Pappus coroniform, of small scales. 

Distribution. The Philippines, Taiwan, S. Japan, and 
China, where it is also widely cultivated. - Monotypic. 

Crossostephium chinense is a former species of Artemisia of 
uncertain systematic position. The rounded capitula with 
their pubescent outer involucral bracts recall many species of 
Artemisia. It appears from inflorescence, floret and pollen 
structure that it is related to the Artemisia group of genera. It 
differs by its weakly ribbed cypselas, furnished with a distinct 
coroniform pappus. On the basis of these characters a pos- 
sible relationship to the North American Artemisia califor- 
nica has been suggested by Gray (1884) and Rydberg (1916). 
Rydberg even transferred some species of Artemisia and 
relatives of Crossostephium on the basis of ribbed cypselas 
being present. However, no pappus is present and the 
presumed relationship was questioned by Hall & Clements 
(1923). 

43. ARTEMISIA L., Sp. pL: 845 (1753). Type 
species: A. vulgaris L. - Oligosporus Cass, 
(including Artemisiella A. Ghafoor) 

Annual and perennial herbs, half-shrubs or shrubs. Leaves 
alternate, variously lobed or dissected, rarely entire. Capitula 
disciform; inflorescence usually a long panicle but sometimes 
much reduced and racemose, spiciform or subglobose. 
Receptacle flat to conical, epaleate, sometimes pilose. Outer 
female florets usually tapering above, with 2-4 teeth, or 
truncate, commonly oblique at orifice. Central florets her- 
maphrodite and fertile or female-sterile and functionally 
male; corolla 5-lobed, yellow or sometimes purplish. Apical 
anther appendage lanceolate-linear to subulate. Cypselas 
obovoid, thin-walled, with or without rows of myxogenic 
cells, usually glabrous but occasionally hairy. Pappus absent. 

Distribution. Predominantly N. hemisphere but with a few 
species also from S. America, Africa S. of Sahara and the 
Hawaiian Islands. Most species in temperate Eurasia and W. 
N. America. - 388 spp. 

Artemisia is the largest genus of the Anthemideae (see 
Krasheninnikov, 1946). Because there are so many species 



different authors have made numerous attempts to divide it 
up in some way. Problems have arisen because two of the 
four commonly recognized sections appear not to be mono- 
phyletic and attempts to separate individual genera have been 
carried out for a variety of different reasons, in regional 
isolation, without an appraisal of either all of the characters 
or all of the taxa. 

The division of Artemisia goes back to Tournefort (1700). 
He recognized three genera, Abrotanum, Absinthium, and 
Artemisia. These were based on gross morphological charac- 
ters and general habit. They are not the same groups as those 
recognized today, although the names have been retained at 
sectional or subgeneric level. Linnaeus (1753) united Tourne- 
fort's three genera into one, establishing more or less the 
concept of Artemisia which has been recognized ever since. 
Cassini (1817) established a new genus, Oligosporus, to 
accommodate those species with functionally separate sexes, 
outer female florets and central, functionally male florets 
with fused style-branches and reduced, abortive ovaries. This 
genus corresponds with the present day section (or subgenus) 
Dracunculus. All of the remaining taxa were included in 
Artemisia, Absinthium not being recognized. The next impor- 
tant development was by Besser (1829). Although he never 
completed his monograph, his results were published by De 
CandoUe (1837). Besser established three subdivisions which 
were expanded to four by de CandoUe as follows: 

Sect. Abrotanum {=Artemisia). Capitula heterogamous (dis- 
ciform); outer florets female, fertile; central florets perfect, 
fertile; receptacle glabrous. 

Sect. Absinthium. As Abrotanum but receptacle hairy. 

Sect. Dracunculus. As Abrotanum but central florets female- 
sterile. 

Sect. Seriphidium. Capitula homogamous (discoid); florets all 
perfect, fertile; receptacle glabrous. 

This arrangement has more or less persisted ever since and 
most treatments have fused or separated the different groups. 
Grenier & Godron (1850) amalgamated all four sections into 
one genus, Euartemisia, but Rouy (1903) by contrast raised 
three sections, Seriphidium, Abrotanum and Absinthium to 
the status of subgenera. Later Rydberg (1916) promoted 
Dracunculus to the same rank. Gray (1884) kept sections 
Seriphidium and Dracunculus but united Abrotanum and 
Absinthium into one new section Euartemisia. Hall & Clem- 
ents (1923) attempted the first phylogenetic interpretation of 
the four sections, on the basis of three transformations. These 
were receptacle becoming hairy, loss of female fertility in disc 
florets and a complete reduction of the female florets. The 
three sections Absinthium, Dracunculus and Seriphidium 
were considered to be three coherent groups derived as three 
separate lines from an ancestral Abrotanum. 

The most recent treatments have been by Russian and 
Chinese botanists (Drokhina, 1978; Poljakov, 1961, 1967; 
Wang, 1979; Leonova, 1971, 1980; Korobkov, 1979). Kra- 
scheninnikov (1946) re-sorted Gray's section Euartemisia, 
raising it to subgeneric rank but kept Dracunculus and 
Seriphidium as two separate subgenera. The most radical 
treatment is that of Poljakov (1961). He used the section 
Artemisia s. s. to accommodate distinctive species of the 
'Artemisia vulgaris' -gvoxxp as distinct from section Abro- 
tanum. This, together with sections Abrotanum and Absin- 
thium, comprised a smaller genus Artemisia. Seriphidium and 
Dracunculus {—Oligosporus) were raised to generic rank 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



121 



together with some other small genera {Kaschgaria, Neopal- 
lasia, Turaniphytum and Mausolea). 

The most recent classifications of Artemisia and its allies 
are those of Y. R. Ling (1980ft, 1982, 1984 1988fl, b, 1991ft). 
In his 1982 treatment he recognizes nine sections oi Artemisia 
and considers Abrotanum and Absinthium the primitive ones, 
from where the others, as well as a number of related genera 
have evolved. It is worth noting that Ling considers 
Seriphidium as a separate genus. Ling (1984) divides the 
genus into two subgenera, Artemisia and Dracunculus, and 
maintains the three traditional sections, Abrotanum, Absin- 
thium and Dracunculus. Seriphidium is kept as a separate 
genus and a number of new combinations are made concern- 
ing Chinese species. 

Absinthium has been variably maintained as a section, sunk 
into subgenus Artemisia (i. e. Abrotanum), or raised to 
subgeneric rank. The main distinguishing feature, and in fact 
the only criterion used for recognition, is the presence of a 
ring of receptacular hairs around the base of each flower. 
However, even Gray (1884) noted that in certain species 
there is only a partial presence, or indeed, a complete 
absence of this character. Poljakov (1961) maintains that it is 
incorrect to sink Absinthium because most species do in fact 
have the character. He notes also that many species of 
Absinthium have a dense, woolly pubescence of white silky 
hairs. As far as we can judge, certain species of different 
sections are artificially separated by this character. Further- 
more, there are those taxa with a dense indumentum but 
without receptacular hairs, and there are those without a silky 
indumentum but with receptacular hairs. The problem is left 
to an internal generic study beyond the scope of this work. 

The section or subgenus Dracunculus is distinguished by 
the fact that the pistil of the central florets is abortive and 
they are therefore functionally male. The central ovaries are 
uniformly sterile and very reduced. This condition is found 
also in Mausolea and Picrothamnus . We agree with Hall & 
Clements (1923) that Dracunculus is a monophyletic group, 
but only so if Mausolea and Picrothamnus are included. 
Future work on generic delimitation of Artemisia s. 1. will 
probably result in Dracunculus being removed from Artemi- 
sia. As a separate genus it will be named Oligosporus, 
following Cassini (1817) and Poljakov (1961). (Dracunculus 
Miller applies to plants of Araceae.) The problem of Mauso- 
lea and Picrothamnus and their sister group, Oligosporus as a 
whole or only a part of it, also has to be considered. Many 
species are involved and several new combinations necessary. 
For these reasons we have for the time being provisionally 
retained Oligosporus as an infrageneric taxon Dracunculus 
within Artemisia. 

Seriphidium, commonly treated as a section or subgenus of 
Artemisia, is recognized here as a separate genus, following 
Poljakov (1961) and Y. R. Ling (1982, 1984). 

Neopallasia and Turaniphytum are two small generic segre- 
gates established by Poljakov (1955, 1961). Apparently they 
have their sister group(s) within Artemisia, paraphyletic as 
presently circumscribed. A proper generic circumscription of 
Artemisia, considering also these segregate genera as well as 
Oligosporus (Artemisia sect. Dracunculus), Picrothamnus 
and Mausolea, is a major task considering the numerous 
species involved. Work on this problem has been undertaken 
by Yeou-Ruenn Ling at The Natural History Museum, Lon- 
don and Institute Sinica, Guangzhou. 

The list of species is compiled from the major floras, with 
recently described species added. From those areas, notably 



China, without recent floristic accounts the list must naturally 
be taken as rather preliminary. In those cases where major 
floras disagree on synonymy, the more recent treatments 
have generally been followed. 

M. abaensis Y. R. Ling & S. Y. Zhao. China. 

A. abrotanum L. Eurasia, widely cultivated and introduced, 

also in N. America. 
A. absinthium L. Eurasia and N. Africa, widely introduced, 

also in N. America. 
A. abyssinica Schultz-Bip. Saudi Arabia. 
A. adamsii Besser. S. Siberia, Mongolia, China. 
* A. afghanica Rech. f. Afghanistan. 
A. afra Jacq. Africa S. of Sahara. 
*A. aksaiensis Y. R. Ling. China. 
A. alaskana Rydb. N. America in Alaska. 
A. albicerata H. Kraschen. C. Asia. 
A. aleutica Hulten. Aleutian Islands. 
*A. altaiensis H. Kraschen. S. Siberia, Mongolia. 
*A. amygdalina Decne. Himalayas. 
A. andersiana Podl. Afghanistan. 
A. anethifolia G. Weber in Stechm. E. Siberia, Mongoha, 

China. 
A. anethoides Mattf. China. 
*A. angustissima Nakai. China, Japan. 
A. annua L. Eurasia, widespread and introduced, also in N. 

America. 
A. anomala S. Moore. China. 
*A. aquatica Lour. China. 
A. arborescens L. S. Europe, Turkey, Middle East, N. Africa 

from Libya to Morocco. 
A. arctica Less. Siberia, Japan, W. North America. 
*A. arctisibirica Korobkov. Siberia. 
A. argyi A. Leveille & Vaniot. Far East, Mongoha, China, 

Korea. 
*A. argyrophylla Ledeb. Mongolia, China. 
A. armeniaca Lam. SE European Russia, Turkey, Iran, S. 

Siberia. 
A. aschurbajewii C. Winkler. C. Asia. 
A. atlantica Cosson & Durieu. N. Africa in Tunisia, Algeria 

and Morocco. 
A. atrata Lam. C. Europe. 
A. atrovirens Hand.-Mazz. China. 
A. aucheri Boiss. Iran, Pakistan. 
*A. aurata V. Komarov. Far East, China, Korea. 
A. australis Less. Hawaii Islands. 
A. austriaca Jacq. E. and E.C. Europe, Turkey, Iran, 

Afghanistan, C. Asia, W. Siberia, Far East, China. 
A. austro-himalayensis (Y. R. Ling & H. S. Puri.) Y. R. Ling 

& H. S. Puri. N. India. 
*A. austro-yunnanensis Ling & Y. R. Ling. China. 
*A. avarica Minat. Caucasus. 

*A. baimaensis Y. R. Ling & Z. C. Z. Y. Zhuo. China. 
*A. banihalensis Kaul & Bakshi. India. 
*A. bargusinensis Sprengel. E. Russia, Siberia. 
*A. bejdemaniae Leonova. Siberia. 
A. biennis Willd. Eurasia, widespread and widely introduced, 

also in N. America. 
*A. blepharolepis Bunge. Mongolia, China. 
A. borealis Pallas - Note: The delimitation towards A. 

campestris is unclear. In Flora europaea (Tutin et al., 1976) 

this species is treated as a subspecies oi A. campestris. N. 

Europe, Siberia, Mongolia, China, N. America. 
A. borealo-siamensis Y. R. Ling. N. Thailand 



122 



K. BREMER AND C. J. HUMPHRIES 



*A. brachyloba Franchet. China. 

*A. brachyphylla Kitam. China, Korea. 

*A. brevis Pampan. China. 

*A. burmanica Pampan. China. 

A. caespitosa Ledeb. S. Siberia, Mongolia, China. 

A. califomica Less. United States in Cahfornia and Mexico in 
Baja Cahfornia. 

*A. calophylla Pampan. China. 

A. campbellii Hook. f. & Thomson. Himalayas, China in 
Tibet. 

A. campestris L. - Note: See note under A. borealis. Wide- 
spread in Eurasia, N. America, and N. Africa. 

A. camphorata Villars. Europe. 

A. canariensis (Besser) Less. Canary Islands. 

*A. cannabifoHa A. Leveille. China. 

A. cantabrica (Lainz) Lainz. SW Europe in Spain. 

A. capillaris Thunb. Far East, China, Japan, SE Asia in 
Malaya. 

A. carruthii Wood. W. United States. 

A. caruifolia Buch.-Ham. in Roxb. Himalayas. 

*A. cashimirica Kaul & Bakshi. India. 

A. caucasica Willd. S. European Russia, Caucasus, Turkey. 

A. chamaemelifolia Villars. C. and SW Europe, Caucasus, 
Turkey, Iran. 

*A. chiajeana Kunze. Iran. 

*A. chiarugii Pampan. China. 

*A. chienshanica Ling & W. W. Wang. China. 

*A. chingii Pampan. China. 

A. chitachensis Cosson ex Battand. & Trabut. N. Africa. 

*A. chrysolepis Kitagawa. China. 

*A. conaensis Ling & Y. R. Ling. China. 

*A. congesta Kitam. Japan. 

A. copa Philippi. S. America (Argentina & Chile). 

*A. coracina W. W. Wang. China. 

A. crithmifolia L. Europe. 

*A. cuspidata H. Kraschen. E. Siberia. 

*A. daghestanica H. Kraschen. & Pors. Caucasus and adja- 
cent parts of Russia (Daghestan). 

*A. dahurica (Turcz.) Polj. China. 

*A. dalai-lamae H. Kraschen. China in Tibet. 

*A. demissa H. Kraschen. C. Asia, China. 

*A. densifolia Filat. Algeria. 

*A. depauperata H. Kraschen. Mongolia. 

A. desertorum Sprengel. Far East, E. Siberia, Mongolia, 
China. 

*A. dimoana Popov. SW & C. Asia. 

A. disjuncta H. Kraschen. Mongolia, China. 

*A. divaricata (Pampan.) Pampan. Mongolia, China. 

M. diversa Diels. China. 

*A. dolichocephala Pampan. Himalayas. 

A. douglasiana Besser in Hook. f. W. United States. 

*A. dracunculiformis H. Kraschen. Arctic Siberia. 

A. dracunculus L. {A. glauca Pallas ex Willd.). Eurasia, 
widely cultivated and introduced, also in N. America. 

A. dubia Wallich ex Besser. Himalayas, China. 

*A. dudinensis V. P. Amel'chenko. Siberia. 

*A. duthreuil-de-rhinsi H. Kraschen. China in Tibet. 

M. edgeworthii Balakr. China. 

A. eldarica Rzazade. Russia. 

*A. elegantissima Pampan. W. Himalayas. 

A. emeiensis (Chang) Y. R. Ling. China. 

A. eriantha Ten. C. and SW Europe. 

*A. eriocephala Pampan. W. Himalayas. 

A. eriopoda Bunge. China. 



*A. erlangshanensis Ling & Y. R. Ling. China. 

*A. faurieri Nakai. Korea, Japan. 

A. filifolia Torrey. W. United States. 

A. filiformilobulata Y. R. Ling & H. S. Puri. India. 

A. flaccida Hand.-Mazz. China. 

*A. flahaultii Emb. & Maire. N. Africa in Morocco. 

A. flava Jurtzev. Siberia. 

*A. flavifolia Gilli. Afghanistan. 

*A. forrestii W. Smith. China. 

A. franserioides Greene. W. N. America. 

A. freyniana (Pampan.) H. Kraschen. Far East, Mongolia, 

China, Korea. 
A. frigida Willd. SE Russia, C. Asia, Siberia, Mongolia, 

China, N. America. 
*A. frigidioides H. C. Fu & Z. Y. Zhu. China. 
*A. fukudo Makino. Korea, Japan. 
*A. fulgens Pampan. China. 
A. f areata M. Bieb. E. Siberia, N. America in Alaska and 

Canada. 
*A. gabriellae Braun-Blanquet. SW Europe. 
A. gangsuensis Ling & Y. R. Ling. China. 
A. genipi G. Weber in Stechm. C. Europe. 
*A. gilveseens Miq. China, Japan. 
A. giraldii Pampan. China. 
*A. glabella KareHn & Kir. China, C. Asia, W. Siberia, 

Mongolia. 
A. glacialis L. C. Europe. 
*A. globosa H. Kraschen. Mongolia, China. 
*A. globosoides Ling & Y. R. Ling. China. 
A. globularia Cham, ex Besser. E. Siberia, N. America in 

Alaska. 
A. glomerata Ledeb. Arctic and E. Siberia, Japan, N. 

America in Alaska and Canada (Yukon). 
A. gmelinii G. Weber in Stechm. Himalayas, C. Asia, 

Siberia, Far East, Mongolia, China, Korea, Japan. 
A. gongshanensis Y. R. Ling & Humphries. China. 
*A. gorgonum Webb in Hook. Cape Verde Islands. 
A. granatensis Boiss. SW Europe in Spain. 
*A. graveolens Minat. Caucasus. 
*A. gyangzeensis Ling & Y. R. Ling. China. 
*A. gyitangensis Ling & Y. R. Ling. China. 
*A. haichowensis Chang. China. 
A. hallaisanensis Nakai. Korea. 
M. halodendron Turcz. ex Besser. E. Siberia, Mongolia, 

China. 
*A. hancei (Pampan.) Ling & Y. R. Ling. China, Indo- 

China. 
*A. haussknechtii Boiss. Turkey, Iraq, Iran. 
A. hedinii Ostenf. & Pauls in Hedin. China in Tibet. 
*A. henriettae H. Kraschen. Arctic Siberia. 
*A. hillebrandii Skottsb. Hawaii Islands. 
*A. hippolytii Butkov. Russia. 
A. hispanica Lam. Spain. 

A. hololeuca M. Bieb. ex Besser. S. European Russia. 
A. hulteniana Vorosch. Aleutian Islands. 
A. hultenii Maksimova. Far East. 
A. idilongensis Y. R. Ling. China. 
*A. ifranensis J. Didier. N. Africa in Morocco. 
*A. igniaria Maxim. China. 
*A. implicata Leonova. Mongolia. 
*A. imponens Pampan. China. 
A. incana (L.) Druce. Turkey, Caucasus and adjacent parts 

of Russia (Daghestan), Iraq, Iran. 
A. incisa Pampan. {A. nuristanica Kitam.). Himalayas. 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



123 



A. indica Willd. Himalayas, China, Taiwan, Japan. 

A. insipida Villars. C. Europe in France. 

A. insulana H. Kraschen. E. Siberia in Bering Island. 

A. integrifolia L. Siberia, Far East, Mongolia, China, Korea. 

*A. intramongolica H. C. Fu & Z. Y. Zhu. China. 

A. jacutica Drob. E. Siberia. 

A. japonica Thunb. Afghanistan, Pakistan, Far East, China, 

Korea, Japan, Taiwan. 
A. javanica Pampan. Indonesia. 
*A. jaxatica Polj. C. Asia. 
A. jilongensis Y. R. Ling & Humphries. China. 
A. judaica L. Middle East and N. Africa in Egypt, Libya and 

Algeria. 
M. kabylica Chabert. N. Africa in Algeria. 
M. kanashiroi Kitam. N. China. 
*A. kangmasensis Ling & Y. R. Ling. China. 
*A. karavajevii Leonova. Siberia. 
A. kauaiensis (Skottsb.) Skottsb. Hawaii Islands. 
A. kawakamii Hayata. Taiwan. 
A. keiskeana Miq. Far East, China, Korea, Japan. 
A. kelleri H. Kraschen. C. Asia. 
A. kitadakensis Hara & Kitam. Japan. 
*A. klementzae H. Kraschen. ex Leonova. Mongolia. 
A. klotzschiana Besser. N. America in Mexico. 
A. koidzumii Nakai. E. Siberia, Far East, Japan. 
*A. komarovii Polj. Far East. 
*A. kulbadica Boiss. & Buhse. Iran, C. Asia. 
*A. kumykorum Minat. Caucasus. 
A. kuschakewiczii Winkler. C. Asia. 
A. laciniatiformis V. Komarov. E. Siberia, Far East, N. 

America in Alaska. 
A. lactiflora Wallich ex DC. China, Taiwan. 
A. lagocephala (Fischer ex Besser) DC. Siberia, Far East, 

China. 
A. lagopus Fischer ex Besser. E. Siberia, Far East. 
A. lamprocaulos Rech. f. Iran. 
A. lancea Vaniot {A. feddei Levi. & Vaniot). China, Korea, 

Japan. 
A. latifolia Ledeb. Russia, C. Asia, Siberia, Mongolia, 

China. 
A. lavandulifolia DC. Far East, China, Korea. 
*A. lavei Kostel. China. 
A. ledebouriana Besser. E. Siberia. 
*A. leontopodioides Fischer ex Besser. E. Siberia, Aleutian 

Islands. 
A. leptophylla D. Don. Himalayas (Nepal). 
A. leucophylla (Turcz. ex Besser) C. B. Clarke. S. Siberia, 

MongoHa, China. 
*A. limosa Koidz. Far East. 

M. limprichtii (Pampan.) Ling & Y. R. Ling. China. 
*A. lipskyi Polj. C. Asia. 
*A. littoricola Kitam. Far East, Japan. 
A. longifolia Nutt. Canada and W. United States. 
A. ludoviciana Nutt. Canada and W. United States. 
*A. macilenta (Maxim.) H. Kraschen. Far East, China. 
M. maciravae Hutch. & Dalziel. Africa in Sahara. 
A. macrantha Ledeb. European Russia, Siberia, Mongoha, 

China. 
A. macrocephala Jacq. Iran, Afghanistan, Himalayas, S. 

Siberia, Far East, Mongolia, China. 
*A. macrorhiza Turcz. E. Siberia. 
A. magellanica Schultz-Bip. S. America in Chile. 
A. mairei A. Leveille. China. 
*A. manshurica (V. Komarov) V. Komarov. Siberia, China. 



A. maroccana Cosson. Morocco. 

*A. marschalliana Sprengel. China. 

*A. martjanovii H. Kraschen. ex Polj. E. Siberia. 

A. mattfeldii Pampan. China. 

A. mauiensis (A. Gray) Skottsb. Hawaii Islands. 

A. maximovicziana (F. Schum.) H. Kraschen. ex Polj. Far 
East. 

*A. medioxima H. Kraschen. ex Polj. Far East, N. China. 

A. melanolepis Boiss. & Kotschy. Iran. 

A. mesatlantica Maire. N. Africa in Morocco. 

A. michauxiana Besser. W. North America. 

A. minor Jacq. in Besser. Himalayas, China in Tibet. 

*A. molinieri Quezel, Barbero & R. Loisel. SW Europe in 
France. 

*A. molluccana Roxb. SE Asia in the Moluccas. 

A. momiyamae Kitam. Japan. 

A. mongolica (Fischer ex Besser) Nakai. China. 

A. monophylla Kitam. Japan. 

A. monosperma Del. Turkey, Middle East, N. Africa in 
Egypt and Libya. 

A. montana Pampan. Far East, China, Japan. 

*A. montevidensis Sprengel. S. America in Argentina. 

*A. moorcroftiana Wallich ex DC. China. 

A. morrisonensis Hayata. Taiwan. 

*A. multisecta Leonova. C. Asia. 

A. mutellina Villars. C. & S. Europe. 

M. myriantha WaUich ex DC. China, Himalyas, Burma. 

M. nakaii Pampan. China, Korea. 

*A. nanshanica H. Kraschen. China in Tibet. 

*A. neglecta Leonova. C. Asia. 

*A. negrei Ouyahya. Morocco. 

*A. nesiotica Raven. W. United States. 

A. niitakayamensis Hayata. Taiwan. 

A. nilagirica (C. B. Clarke) Pampan. India, Burma. 

A. nitida Bertol. C. Europe. 

*A. nivalis Braun-Blanquet. C. Europe in Switzerland. 

*A. nortonii Pampan. China in Tibet. 

A. norvegica Fries. N. Europe, Arctic America. 

*A. nujianensis (Ling & Y. R. Ling) Y. R. Ling. China. 

A. obscura Pampan. MongoHa, China. 

A. obtusiloba Ledeb. C. Asia, S. Siberia, Mongolia. 

*A. occidentali-sichuansensis Y. R. Ling. China. 

A. occidentali-sinensis Y. R. Ling. China. 

A. occidentali-yunnanensis Ling & Y. R. Ling. China. 

A. oelandica (Besser) V. Komarov. N. Europe in Sweden. 

*A. olchonensis Leonova. Siberia. 

*A. olgensis (Vorobiev) Vorosch. Russia. 

A. oligocarpa Hayata. Taiwan. 

A. opulenta Pampan. N. Japan, E. Russia. 

*A. oranensis Filat. Algeria. 

*A. ordosica H. Kraschen. China. 

*A. orientalis (Pampan.) Ling & Y. R. Ling. China. 

A. orientali-hengduangensis Ling & Y. R. Ling. China. 

A. orientali-xizangensis Y. R. Ling & Humphries. China. 

A. orientali-yunnanensis Y. R. Ling. China. 

*A. orthobotrys Kitagawa. China. 

*A. oxycephala Kitagawa. China. 

*A. packardiae Grimes & Ertter. W. United States. 

*A. pallens Wallich ex Besser. India. 

A. palustris L. S. Siberia, Far East, Mongolia, China. Mon- 
golia, China, Korea. 

A. pancicii (Janka) Ronniger. E.C. Europe. 

M. pannosa H. Kraschen. Far East. 

A. papposa Blake & Cronq. W. United States in Idaho. 



124 



K. BREMER AND C. J. HUMPHRIES 



A. parry i A. Gray. W. United States. 

A. parviflora Buch.-Ham. ex Roxb. China. 

A. pattersonii A. Gray. W. United States. 

*A. pedatifida Nutt. W. and C. United States. 

A. pedunculosa Miq. Japan. 

M. pengchuoensis Y. R. Ling & S. Y. Zhao. China. 

A. persica Boiss. Iran, Afghanistan, Himalayas, C. Asia. 

*A. pewzowii Winkler. China in Tibet. 

*A. phaeolepis H. Kraschen. S. Siberia, Mongolia, China. 

*A. phyllobotrys (Hand.-Mazz.) Ling & Y. R. Ling. China. 

*A. polybotryoidea Y. R. Ling. China. 

A. pontica L. C. and E. Europe, W. Siberia, introduced in N. 

America. 
M. porteri Cronq. W. United States in Wyoming. 
*A. praticola Klokov. Ukraine. 
*A. prattii (Pampan.) Ling & Y. R. Ling. China. 
A. princeps Pampan. China, Korea, Japan, Taiwan. 
*A. przewalskii H. Kraschen. China. 
A. pseudopontica Schur. E. Europe. 
*A. pubescens Ledeb. China. 
*A. punctigera H. Kraschen. Far East, European Russia, E. 

and W. Siberia, Mongolia, N. America. 
*A. pycnorhiza Ledeb. C. Asia, S. Siberia, MongoUa. 
*A. quinlingensis Ling & Y. R. Ling. China. 
*A. quinqueloba Trautv. C. Asia. 
*A. ramosa C. Smith. Canary Islands. 
*A. rehan Chiov. Africa in Ethiopia. 
*A. remotiloba H. Kraschen. ex Polj. E. Siberia. 
A. reptans C. Smith ex Link. SW Europe in Spain, N. Africa 

in Morocco, Canary Islands. 
*A. robusta (Pampan.) Ling & Y. R. Ling. China. 
*A. rosthornii Pampan. China. 
A. roxburghiana Besser. Afghansitan, Pakistan, Himalayas, 

China. 
M. rubripes Nakai. Far East, MongoHa, China, Korea, 

Japan. 
A. rupestris L. N. Europe, C. Asia, W. Siberia, Mongolia, 

China, N. America in Canada. 
A. rutifolia Stephen ex Sprengel. Iran, Afghanistan, Himala- 
yas, C. Asia, Siberia, Mongolia, China. 
A. sacrorum Ledeb. China, Korea, Japan, Himalayas, C. 

Asia, Afghanistan. 
*A. saitoana Kitam. Far East, Korea. 
A. salsoloides Willd. Russia, W. Himalayas, W. Siberia, 

China in Tibet. 
*A. samoiedorum Pampan. Arctic Siberia. 
A. santolinifoUa Turcz. ex H. Kraschen. Afghanistan, China 

in Tibet, European Russia, C. Asia, S. Siberia, Mongolia, 

Pakistan. 
*A. saposhnikovii H. Kraschen. ex Polj. C. Asia. 
A. schimperi Schultz-Bip. ex Schweinf. Africa in Ethiopia. 
*A. schischkiniiW. Kraschen. Mongolia. 
A. schmidtiana Maxim. Far East, Japan. 
A. scoparia Waldst. & Kit. C. and E. Europe, Turkey, 

Middle East, Iran, Himalayas, C. Asia, Siberia, Mongolia, 

China, Japan, N. Africa in Egypt. 
A. scopulorum A. Gray. W. United States. 
A. selengensis Turcz. ex Besser. E. Siberia, Far East, Mongo- 
lia, China. 
A. senjavinensis Besser. E. Siberia, N. America in Alaska. 
A. sericea G. Weber in Stechm. Russia, Siberia, Mongolia, 

China. 
A. serrata Nutt. W. United States. 
M. serreana Pampan. China. 



M. shangnanensis Ling & Y. R. Ling. China. 

*A. shennongjaensis Ling & Y. R. Ling. China. 

*A. sichuanensis Ling & Y. R. Ling. China. 

A. sieversiana Ehrh. in Willd. European Russia, Himalayas, 
C. Asia, Siberia, Far East, China. 

*A. simulans Pampan. China. 

A. sinanensis Yabe. Japan. 

A. sinensis (Pamp.) Ling & Y. R. Ling. China. 

A. smithii Mattf. China. 

*A. somai Hayata. Taiwan. 

A. songarica Schrenk. C. Asia, China. 

*A. speciosa (Pampan.) Ling & Y. R. Ling. China. 

*A. sphaerocephala H. Kraschen. Mongolia, China. 

A. splendens Willd. Turkey, Caucasus, Iraq, Iran. 

A. stelleriana Besser. Far East, China, introduced in N. 
Europe and N. America, Japan. 

A. stenophylla Kitam. China, Korea, E. Russia. 

A. stipularis Urb. & Ekman. Haiti. 

A. stolonifera (Maxim.) V. Komarov. Far East, China, 
Japan. 

A. stracheyi Hook. f. & Thomson ex C. B. Clarke (Note 
added in proof. This taxon was recently removed by 
Ghafoor (1992) and described as a monotypic genus under 
the name Artemisiella stracheyii (Hook. f. & Thomson ex 
C. B. Clarke) Ghafoor). Himalayas, China in Tibet. 

A. striata Edgew. Iran, Himalayas, China in Tibet. 

*A. subulata Nakai. Soviet Far East, China, Korea. 

A. subviscosa Turcz. E. Siberia. 

*A. succulenta Ledeb. C. Asia, W. Siberia. 

*A. succulentoides Ling & Y. R. Ling. China. 

A. suksdorfii Piper. W. Canada and United States. 

*A. superba Pampan. Mongolia. 

A. swatensis Podl. Pakistan. 

A. sylvatica Maxim. Far East, Mongolia, N. China. 

*A. tafelii Mattf. China in Tibet. 

A. taibaishanensis Y. R. Ling & Humphries. China. 

*A. tainingensis Hand.-Mazz. China. 

A. tanacetifolia L. China, N. Korea, C. and W. Russia, 
Europe, N. America. 

A. tangutica Pampan. China. 

A. tenuifolia Y. R. Ling & H. S. Puri. China. 

A. thellungiana Pampan. SW China, N. India, Sikkim. 

A. tilesii Ledeb. Arctic Eurasia and Arctic N. America. 

A. tomentella Trautv. C. Asia. 

A. tournefortiana Reichenb. Turkey, Caucasus, Iran, 
Afghanistan, Himalayas, C. Asia, Mongolia, China. 

*A. transbaicalensis Leonova. Siberia. 

*A. trautv etteriana Besser. S. European Russia. 

A. tridactyla Hand.-Mazz. China. 

*A. triniana Besser. Arctic Siberia. 

A. tschernieviana Besser. E. Europe, C. Asia, China. 

*A. tsugitakaensis (Kitam.) Ling & Y. R. Ling. China. 

*A. tsuneoi Tatewaki & Kitam. Japan. 

*A. tukuchaensis Kitam. Himalayas (Nepal). 

A. tyitangensis Ling & Y. R. Ling. China. 

A. unalaskensis Rydb. Far East, N. America in Alaska. 

M. ussuriensis Polj. Far East. 

*A. velutina Pampan. China. 

*A. verbenacea (V. Komarov) Kitagawa. China. 

A. verlotorum Lamotte. China, Himalayas, Malaya, natural- 
ized in W. and C. Europe and in N. Africa, S. America. 

A. vestita Wallich ex Besser. W. Himalayas. 

A. vexans Pampan. China, Himalayas. 

*A. viridisquama Kitam. China. 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



125 



*A. viridissima (V. Komarov) Pampan. China. 

M. viscida (Mattf.) Pampan. Himalayas, China. 

M. viscidissima Ling & Y. R. Ling. China. 

A. vulgaris L. Widespread in Eurasia and N. America, also in 

N. Africa, widely introduced e. g. in AustraUa. 
*A. waltonii J. R. Drumm. ex Pampan. China in Tibet. 
*A. wellbyi Hemsley & Pears. China, Himalayas. 
*A. wudanica Liou & W. Wang. China. 
*A. xanthochloa H. Kraschen. Mongolia, China. 
*A. xerophytica H. Kraschen. Mongolia, China. 
M. xigazeensis Ling & Y. R. Ling. China. 
M. yadongensis Ling & Y. R. Ling. China. 
A. yongii Y. R. Ling. China. 
*A. younghusbandii J. R. Drumm. China in Tibet. 
A. yunnanensis Jeffrey ex Diels. China. 
M. zayuensis Ling & Y. R. Ling. China. 
A. zhongdianensis Y. R. Ling. China. 

44. NEOPALLASIA Polj. in Bot. Mater. Gerb. hot. 
Inst. V. A. Komarova 17: 429 (1955). Type species: 
N. pectinata (Pallas) Polj . 

Annual or biennial herbs. Leaves alternate, pectinate- 
pinnatisect with filiform, apically somewhat swollen and 
mucronulate lobes. Capitula rather small and rounded in a 
narrow spiciform panicle, disciform. Receptacle narrowly 
conical, epaleate. Outer female florets narrowly tubular, 
without teeth. Central florets of two kinds; outer perfect, 
inner completely sterile with reduced ovaries. Apical anther 
appendage ovoid-lanceolate and acuminate. Cypselas 
arranged around the base of the receptacle, oblong-obovoid, 
somewhat compressed or triquetrous, thin-walled, with many 
rows of myxogenic cells. Pappus absent. 

Distribution. C. Asia, S. Siberia, MongoUa and China. - 3 
spp. 

Poljakov (1955) distinguished Neopallasia from Artemisia by 
the characteristic pectinate leaves, the apically truncate (with- 
out teeth) outer female florets, the presence of completely 
sterile central florets (in addition to perfect ones) situated at 
the apex of a narrowly conical receptacle, the ovoid- 
lanceolate and attenuate anther appendages and the rosette- 
shaped arrangement of the cypselas around the receptacle. 
These characters are autapomorphies, though the shape of 
the anther appendages is hardly spectacular considering the 
variation present within the subtribe, and the cypsela 
arrangement follows from the sex distribution within the 
head, fertile florets being restricted to the outer part or the 
base of the conical receptacle. 

The immediate relatives or the sister group of Neopallasia 
is not easy to identify. Poljakov suggested that the genus is 
related to Artemisia sect. Dracunculus because some of the 
central hermaphrodite florets are sterile. It does seem clear 
that Neopallasia has its sister group within a presently para- 
phyletic Artemisia, possibly within sect. Dracunculus as sug- 
gested by Poljakov. 

Y. R. Ling (1980fl) has recently added two Chinese species. 
The material of those were formerly considered part of N. 
pectinata s. 1. 

A^. pectinata (Pallas) Polj . 

N. tibetica Y. R. Ling 

*N. yunnanensis (Pampan.) Y. R. Ling 



45. TURANIPHYTUM Polj. in Komarov, Fl. URSS 
26: 880 (1961). Type species: T. eranthemum 
(Bunge)Polj. 

Perennial herbs, somewhat woody at the base. Leaves alter- 
nate to rosulate, pinnatisect. Capitula disciform; inflores- 
cence a spike of glomerules with densely congested capitula, 
or rarely capitula solitary in interrupted, partly congested 
spikes. Receptacle convex to hemispherical, epaleate. Outer 
female florets subtended by scaphoid inner involucral bracts, 
unequally crenate at the apex. Central florets 5-lobed, api- 
cally with long rigid somewhat reddish hairs at the apex. 
Apical anther appendage lanceolate, acuminate. Cypselas 
obliquely oblong-obovoid, thin-walled, with rows of myxo- 
genic cells. Pappus absent. 

Distribution. C. Asia. - 2 spp. 

Turaniphytum was distinguished from Artemisia mainly 
because of the peculiar inflorescence, with the capitula aggre- 
gated into glomerules and arranged in long spikes. These are 
presumably transformed paniculate inflorescences of the 
common Artemisia type. Turaniphytum also has scaphoid 
inner involucral bracts, subtending the outer female florets. 
The immediate relatives of Turaniphytum are unknown. It 
may have its sister group within Artemisia. 

T. codringtonii (Rech. f.) Polj. (T. kopetdaghense Polj.). 

Afghanistan. 
T. eranthemum (Bunge) Polj. 

46. MAUSOLEA Polj. in Trudy Inst. Bot. Alma-Ata 
11: 170 (1961). Type species: M. eriocarpa (Bunge) 
Polj. 

A virgate shrub. Leaves alternate, few-lobed or entire. 
Capitula small and subglobose, rather few and more or less 
sessile in a reduced panicle, disciform. Receptacle epaleate. 
Outer female florets without corolla; style-branches dilated, 
lanceolate, flat, acute. Central florets 5-lobed, apically with 
bifurcate hairs, hermaphrodite and female-sterile; ovaries 
reduced and style-branches fused. Apical anther appendage 
narrowly lanceolate-linear. Cypselas obovoid, densely pilose. 
Pappus absent. 

Distribution. Iran, Afghanistan, and C. Asia. - Mono- 
typic. 

Mausolea was separated from Artemisia mainly because of 
the corollaless marginal flowers. The styles of the marginal 
flowers are also further modified compared to those of 
Artemisia, being wider and lanceolate. It is probably related 
to Picrothamnus as discussed under that genus. The cypsela 
hairs are straighter and less cobwebby in Mausolea compared 
to Picrothamnus. 

47. PICROTHAMNUS Nutt. in Trans. Amer. Philippi 
Soc. II, 7: 417 (1841). -Type species: P. desertorum 
Nutt. 

A basally much woody shrublet with older branches trans- 
formed into long spines. Leaves alternate, few-lobed. 
Capitula small and subglobose, solitary or few together along 
the branches, almost sessile, disciform. Receptacle epaleate. 
Outer female florets tubular. Central florets 5-lobed, with 
long cobwebby bifurcate hairs, hermaphrodite and female- 
sterile; ovary reduced and style-branches fused. Apical 



126 



K. BREMER AND C. J. HUMPHRIES 



anther appendage lanceolate-linear. Cypselas obovoid, thin- 
walled, densely cobwebby-pilose with bifurcate hairs. Pappus 
absent. 

Distribution. N. America in western United States. - 
Monotypic. 

Picrothamnus was established by Nuttall but reduced by 
Eaton (in Watson, 1871), a classification accepted by most 
later authors. Hence it is generally known as Artemisia 
spinescens D. C. Eaton. Hall & Clements (1923) consider it 
'in all essentials an Artemisia of the section Dracunculus\ 
The spiny habit and the cobwebby-pilose corollas are autapo- 
morphies of Picrothamnus. The cobwebby-pilose cypselas are 
shared with Mausolea, the probable sister group. Together 
they are related to Artemisia sect. Dracunculus because of 
their functionally male central florets with reduced ovaries 
and fused style-branches. Pending a revised generic delimita- 
tion of Artemisia we think these genera should be retained 
rather than sunk in Artemisia. 

7. ACHILLEINAE Bremer & Humphries, 
subtrib. no v. 

Type species: Achillea millefolium L. 

Herbae annuae vel perennes vel suffrutices. Receptaculum 
paleaceum. Corolla flosculorum disci tubo varie saccato et 
incrassato, saltern adaxialiter basi saccato. Cypselae parieti- 
bus tenuibus, plerumquefasciculis vascularibus duobus later- 
alibus interdum etiam fasciculo uno adaxiali vel nonnunquam 
fasciculis 4-5 instructae, interdum compressae, saepe cellulis 
mucilaginis instructae. Pappus nuUus. 

Annual or perennial herbs or shrublets. Leaves variously 
dissected, sometimes vermiform, rarely few-lobed or entire. 
Capitula solitary or corymbose, radiate or discoid. Recep- 
tacle variously shaped, often conical, paleate. Ray floret limb 
white or yellow. Disc corolla 5-lobed; tube variously saccate 
and thickened in fruit, basally saccate at least adaxially. 
Cypselas thin-walled, generally with 2 lateral and with or 
without 1 adaxial strand, sometimes 4— 5-stranded, sometimes 
flattened, often with myxogenic cells. Pappus absent. 

Distribution (Table 16): Eurasia and N. Africa, mainly in 
S. Europe, the Mediterranean and SW Asia, also in N. 
America, some species of Achillea widely introduced also in 
the S. hemisphere and one species of Santolina introduced in 
N. America. - 9 genera, 147 spp. 

Some groupings within this subtribe have been recognized 



earlier. The relationship between Achillea, Anacyclus, and 
Leucocyclus has been pointed out by Humphries (1979) and 
phytochemical investigations have indicated a close relation- 
ship between Chamaemelum and Cladanthus. Both genera 
accumulate similar thiophene derivatives (Greger, 1977). 
Phytochemistry has also contributed to the recognition of 
subtribe Achilleinae. Achillea and Anacyclus, Chamaemelum 
and Cladanthus, as well as Otanthus, all synthesize amides 
(Greger, 1977; Bohlmann et al., 1973). Santolina has a 
different chemistry, plesiomorphic in being similar to Tanace- 
tum, but it is here provisionally accepted in Achilleinae 
because of its adaxially saccate corolla and the paleate 
receptacle. Leucocyclus, Mecomischus, and Rhetinolepis , 
three little-known North African genera, are also included in 
Achilleinae and considered related to Chamaemelum and 
Cladanthus on a number of morphological characters. Meco- 
mischus and Rhetinolepis have not been investigated chemi- 
cally. 

The cladogram is one of nine equally parsimonious cla- 
dograms. They involve rearrangements at the base and 
among the last four genera. In all cladograms and hence also 
in the strict consensus tree, clades Ac4 {Achillea, Anacyclus, 
Leucocyclus), Ac5 {Anacyclus and Leucocyclus), Ac6 {Meco- 
mischus, Chamaemelum, Rhetinolepis, Cladanthus), and Ac8 
{Rhetinolepis and Cladanthus) were present. 

Clades and characters - Fig. 8, Tables 2, 17. 



|=Acl=ij= 48 Santolina 

n= 49 Otanthus 



=:AC24 



50 Achillea 



=Ac3 



Ac 4 



Ac 6 



C3U Acniiiea 
rj= 51 Anacyclus 
— Ac 5^ 52 Leucocyclus 
rj= 53 Mecomischus 



€ 



54 Chamaemelum 



Ac 8 



[[= 55 Rhetinolepis 
J^ 56 Cladanthus 



Fig. 8 Cladogram (of nine possible) of the Achilleinae produced 
by the ie option. Cladogram length = 34, consistency index = 88, 
retention index = 85. 

Clade Acl - subtribe Achilleinae 

45 Receptacle paleate. 

82 Disc corolla tube thickened in fruit. This character is 
variously strongly expressed in the different genera, most 
clearly in Otanthus. It is hardly evident in Achillea, Anacy- 
clus, and Leucocyclus. 



Table 16 General distribution of Achilleinae and genera. x=indigenous, o=introduced. 





N.Am. 


Eur- 
Asia 


C.&E. 

Asia 


SW 

Asia 


S.Eur. 


N.Afr. 


S.Afr. 


Austr. 
N.Zeal. 


S.Am 


Achilleinae 


o 


X 


x 


X 


X 


X 


o 


o 


o 


Santolina 


o 








X 


X 








Otanthus 




X 


x 


X 












Achillea 


o 


X 


x 


X 


X 


X 


o 


o 





Anacyclus 

Leucocyclus 

Mecomischus 




X 
X 






X 


X 
X 








Chamaemelum 




X 






X 


X 








Rhetinolepis 
Cladanthus 










X 


X 

X 









GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 

Table 17 Data matrix for the Achilleinae. 1 = presence, = absence, 
? = missing data or not applicable, p = polymorphic but scored as 
the plesiomorphic condition, a = polymorphic but scored as the 
apomorphic condition. 

nil 11 1 111 111 1 

14505765 4887 38 28351116284358 351471 
53271811 524225199608479175194017401836 

48. Santolina al?allla allll lOOOOOOOOOOpOOOOOOOOOpOOO 

49. Otanthus 01?al?la all 101 11 1 100000000000000000000 

SO.Achillea alaallla aOllOplOpOl 1 lOOpOOOOOOOOOOpOOO 

51. Anacyclus alaal?la aOllOplOOOl 1 1 1 laOOOOOOpOOOOOOO 

52. Leucocyclus alaal?10 aOl 10070001 1 1 1 101 1000000000000 

53. Mecomischus alaal?la alll00?00010000000all?p0000000 
54.Chamaemelumalaan\a alUOplOOOlOOOOOOOalUpOOOOOOO 

55. Rhetinolepis al?al?la alll01?00010000000011?l 111 1000 

56. Cladanthus alOal?la alllOOlOOOlOOOOOOOllU 1 111011 1 

1111 11 1 

167772555556666617 141383 
3605613567905678513246726 

48.Santolina ??????????????????0p00000 

49. Otanthus ??????????????????0000000 

50.Achillea ???00000000000000000pppp0 

51. Anacyclus ???00000000000000000p0000 

52. Leucocyclus ?????00000000000000000000 
53. Mecomischus ?????00000000000000000000 
54.C/iflmaeme'/MAn???000000000000000000000p 
55. Rhetinolepis ???????????????????000000 
56.Cladanthus ???0000000000000000000000 



84 Disc corolla tube basally saccate at least adaxially. 

Ill Pappus absent in ray and disc cypselas. 

Santolina 

2 Plants shrubby. 

35 Capitula discoid. Some Anacyclus and Chamaemelum 
species are also discoid, as well as Otanthus and Rhetinolepis. 

Clade Ac2 

181 Amides present. Leucocyclus, Mecomischus, and Rhetin- 
olepis have not been investigated chemically. 

Otanthus 

9 Plants covered with a dense greyish-white indumentum. 

15 reversed. Leaves not variously deeply lobed or divided, but 
entire or crenulate only. 

29 Capitula densely corymbose. Most species of Achillea also 
have densely corymbose capitula. 

35 Capitula discoid. 

86 Corolla basally copiously swollen and spongy, almost 
enclosing the cypsela especially laterally. 

Clade Ac3 

130 Cypselas with 2 lateral vascular strands, sometimes also 
with 1 adaxial strand. This cypsela vascularization is also 
characteristic of a large part of subtribe Matricariinae, but the 
two groups do not seem closely related. 

Clade Ac4 

58 Ray and disc floret tube dorsiventrally flattened. 

82 reversed. See clade Acl. 



127 

114 Cypselas dorsiventrally flattened. Flattened cypselas occur 
in various subtribes. 

Achillea 

There is no obvious autapomorphy for this large genus. 

Clade Ac5 

117 Cypselas laterally winged. Some genera of the Cotula 
group (Matricariinae) also have winged cypselas. As noted 
above (clade Ac3) the groups are not closely related. 

119 Cypselas with sclerenchymatic lateral wings. 

Anacyclus 

61 Ray floret tube persistent on the cypsela. This character 
occurs also in some genera of subtribe Thaminophyllinae. 

Leucocyclus 

27 Leaves vermiform. Some species of Santolina also have 
vermiform leaves. 

51 reversed. Floral parts without resin canals. 

85 Disc corolla deeply and equally saccate both abaxially and 
adaxially. 

Clade Ac6 

41 Involucral bracts wide, flabelliform. This character 
reverses in the small-headed genus Rhetinolepis. 

139 Cypselas completely covered with rows of myxogenic cells. 

154 Cypselas thin-walled, obovoid to oblanceolate, devoid of 
ribs. The same type of cypselas is characteristic of the 
unrelated Artemisia and allies in subtribe Artemisiinae. 

Mecomischus 

There is no obvious autapomorphy for this little-known 
genus. 

Clade Ac7 

180 Particular thiophene derivatives present. Rhetinolepis has 
not been investigated chemically. 

Chamaemelum 

There is no autapomorphy for Chamaemelum. The genus is 
variable in several characters. 

Clade Ac8 

I Plants annual. Some species of Chamaemelum and other 
genera of the Achilleinae are also annuals. 

7 Plants with branches in whorls below the first capitula. The 
capitula of Rhetinolepis and Cladanthus are very different in 
size and shape, but sessile (following character) and arranged 
similarly. 

34 Capitula sessile along the stems. 

50 Receptacular paleae pilose. This character also occurs in 
some species of Chamaemelum, as well as in the unrelated 
Eriocephalus in subtribe Matricariinae. 

Rhetinolepis 

II Plants with dolabriform hairs. 

35 Capitula discoid. 



128 



K. BREMER AND C. J. HUMPHRIES 



41 reversed. See clade Ac6. 

Cladanthus 

48 Receptacle pilose. Some genera of subtribe Thaminophylli- 
nae also have pilose receptacles. 

52 reversed. Ray floret limb not white, but yellow. 

73 Disc corolla lobes with dorsal appendages. Corolla lobe 
appendages occur also in various genera of subtribes Leucan- 
theminae and Matricariinae. 

116 Disc cypselas laterally flattened. Laterally flattened but 
otherwise quite different cypselas occur in subtribe Chrysan- 
theminae. 

48. SANTOLINA L., Sp. pi.: 842 (1753). Type 
species: S. chamaecyparissus L. 

Shrublets. Leaves alternate, dentate to pinnatifid or pinnati- 
sect, sometimes vermiform. Capitula soUtary, pedunculate, 
discoid. Receptacle convex, paleate; paleae with a central 
resin canal. Corolla 5-lobed, basally saccate around the 
cypsela especially adaxially, with a rather long bent tube and 
a distinct limb. Cypselas 3-5-angled, sometimes with myxo- 
genic cells. Pappus absent. 

Distribution. S. Europe, mainly in Spain but extending to 
Jugoslavia, and N. Africa in Morocco and Algeria, one 
species (5. rosmarinifolia) introduced in N. America. - 8 spp. 

Santolina is provisionally placed in Achilleinae. It differs in 
chemistry from the other genera of this subtribe, containing 
polyacetylenes and other substances similar to those in Tan- 
acetum (Greger, 1977). It may be a derivative of that genus 
rather than the sister group of the genera of Achilleinae. 
Although some efforts have already been made to identify 
taxa based on chromosome numbers (e.g. Arrigoni, 1977) 
Santolina is in need of revision. There are two species, S. 
chamaecyparissus and S. rosmarinifolia, with numerous syn- 
onyms and many variants. A few 'species' have been 
described from North Africa. Apparently they belong under 
the complex European species. 

5. chamaecyparissus L. 

5. elegans Boiss. ex DC. 

*S. insularis (Gennari ex Fiori) Arrig. 

*S. ligustica Arrig. 

*S. marchi Arrig. 

5. oblongifolia Boiss. 

5. rosmarinifolia L. 

S. viscosa Lagasca 

49. OTANTHUS Hoffsgg & Link, Ft. portug. 2: 364 
(1889). Type species: O. maritimus (L.) Hoffsgg & 
Link - Diotis Desf . 

A suffruticose perennial covered with a dense greyish-white 
indumentum. Leaves alternate, entire or crenulate. Capitula 
corymbose, discoid. Receptacle convex, paleate; paleae with 
a central resin canal. Corolla 5-lobed, basally copiously 
swollen and spongy, almost enclosing the cypsela especially 
laterally. Cypselas with 4-5 weak ribs, thin-walled, glandular. 
Pappus absent. Amides present. 

Distribution. Europe, N. Africa, and SW Asia extending 



from Ireland to Caucasus, along sea shores, mainly in the 
Mediterranean. - Monotypic. 

This characteristic species is known also as Diotis candidis- 
sima Desf. The interrelationships of Otanthus have been 
obscure but it seems that it is an autapomorphic member of 
subtribe Achilleinae, sharing the same chemistry as most 
members of the subtribe. The copiously swollen corolla base 
also indicates the same relationship, though the character is 
not so extremely developed in the other genera. 

50. ACHILLEA L., Sp. pl.\ 896 (1753). Type species: 
A. millefolium L. 

Perennial herbs generally with rhizomes. Leaves alternate, 
pinnatisect, lobed or rarely entire. Capitula comparatively 
small, generally corymbose or rarely few together or solitary, 
radiate or rarely discoid. Receptacle flat to convex or conical 
or rarely much elongated, paleate; paleae sometimes with a 
central resin canal. Ray florets female, fertile; limb rather 
short and wide, white or yellow; tube more or less flattened. 
Disc corolla 5-lobed, more or less flattened, basally slightly 
saccate around the cypsela especially adaxially, white, yel- 
low, or pink. Cypselas dorsiventrally flattened, with two 
lateral ribs with vascular strands, a third adaxial vascular 
strand rarely present. Pappus absent. Amides present. 

Distribution. Europe and temperate Asia, some species 
also in N. Africa, a few species, in general A. millefolium (s. 
1.) naturalized in N. America and also in the S. hemisphere; 
most species in SE Europe and SW Asia. - 115 spp. 

Achillea is a large genus, but with respect to floral characters, 
homogeneous and well defined. The cypselas are flattened 
and thin-walled with 2 lateral ribs and the corolla is basally 
slightly saccate around the cypsela (Khandzhyan, 1983). The 
Hst of species is compiled from standard floras. 

A. abrotanoides (Vis.) Vis. SE Europe. 

A. absinthoides Hal. SE Europe (Greece). 

A. acuminata (Ledeb.) Schultz-Bip. E. Siberia, Far East, 
Mongolia, China, Japan. 

A. aegyptiaca L. SE Europe (Greece). 

A. ageratifolia (Smith in Sibth. & Smith) Boiss. SE Europe. 

A. ageratum L. S. Europe and N. Africa in Morocco. 

A. aleppica DC. Turkey, Middle East, Iraq, Iran. 

*A. alpina L. E. Siberia, Far East, Mongolia, China, Himala- 
yas. 

M. ambrosiaca (Boiss. & Heldr.) Boiss. SE Europe 
(Greece). 

A. arabica Kotschy. Middle East. 

M. asiatica Serg. C. Asia, Siberia, Far East, Mongolia, 
China. 

A. asplenifolia Vent. E. Europe. 

A. atrata L. C. Europe. 

A. aucheri Boiss. Iran. 

A. barbeyana Heldr. & Heimerl in Heimerl. SE Europe 
(Greece). 

A. barrelieri (Ten.) Schultz-Bip. S. Europe (Italy). 

A. biebersteinii Afan. {A. micrantha Willd., A. micranthoides 
Klokov). E. Europe, S. European Russia, Turkey, Cauca- 
sus, Middle East, Iran, Afghanistan, C. Asia. 

A. biserrata M. Bieb. Caucasus. 

*A. boissieri (Hausskn.) Boiss. Turkey. 

*A. brachyphylla Boiss. & Hausskn. in Boiss. Turkey. 

*A. bucharica Winkler. C. Asia. 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



129 



A. callichroa Boiss. Iran. 

A. camtschatica Rupr. ex Heimerl. E. Siberia. 

A. cappadocica Hausskn. & Bornm. Turkey. 

A. cartilaginea Ledeb. ex Reichenb. {A. septentrionalis 

(Serg.) Botsch.). E. Europe, Siberia, C. Asia. 
A. chamaemelifolia Pourret. SW Europe. 
A. chrysocoma Friv. SE Europe. 
A. clavennae L. C. and SE Europe. 
A. dypeolata Sibth. & Smith SE Europe. 
A. coarctata Poiret in Lam. SE Europe, Turkey. 
A. collina J. Becker ex Reichenb. C. and SE Europe. 
A. compacta Willd. SW Europe, S. Russia. 
A. conferta DC. Syria, Iraq, Iran. 
A. cretica L. SE Europe, Cyprus, Turkey. 
A. crithmifolla Waldst. & Kit. C. and SE Europe. 
*A. cucullata (Hausskn.) Bornm. Turkey. 
*A. cuneatiloba Boiss. & Buhse. Caucasus, Iran. 
*A. decolorans Schrader. Turkey. 
A. depressa Janka. E. and SE Europe. 
A. distans Waldst. & Kit. ex Willd. C. Europe. 
A. erba-rotta All. C. and SE Europe. 
A. falcata L. Turkey, Middle East, Iraq. 
A. filipendulina Lam. Caucasus, Iran, Afghanistan, C. Asia. 
A. fraasii Schultz-Bip. Turkey, SE Europe. 
A. fragrantissima (Forssk.) Schultz-Bip. N. Africa in Egypt, 

Middle East, Iraq. 
A. gerberi Willd. W. Asia, S. Europe, Russia. 
A. glaberrima Klokov. S. European Russia. 
A. goniocephala Boiss. & Bal. in Boiss. Turkey. 
A. grandifolia Friv. SE Europe, Turkey. 
A. griseo-virens Albov. Caucasus. 
A. gypsicola Huber-Mor. Turkey. 
A. holosericea Sibth. & Smith. SE Europe. 
A. impatiens L. E. Europe (Romania), Siberia, China. 
A. inundata Kondr. in Wissjul. S. European Russia. 
*A. japonica Heimerl. Far East, China, Japan. 
*A. kellalensis Boiss. & Hausskn. in Boiss. {A. haussknechtii 

Boiss.). Iran. 
A. kotschyi Boiss. Turkey. 
*A. latiloba Ledeb. ex Nordm. Caucasus. 
*A. ledebourii Heimerl. S. Siberia, China. 
A. leptophylla M. Bieb. E. Europe, S. European Russia, N. 

Africa in Morocco and Algeria. 
A. ligustica All. S. Europe, N. Africa in Morocco, Algeria 

and Tunisia. 
A. lingulata Waldst. & Kit. SE Europe. 
A. lucana Pign. Italy. 

A. lycaonica Boiss. & Heldr. in Boiss. Turkey. 
*A. macrocephala Rupr. Far East, Japan. 
A. macrophylla L. C. Europe. 
A. magnifica Huber-Mor. Turkey. 
A. maura Humbert. N. Africa in Morocco. 
A. membranacea (Labill.) DC. Turkey, Middle East, Iraq. 
A. millefolium L. {A. lanulosa Nutt., A. sudetica Opiz). 

Widespread in Eurasia and N. America, introduced in 

Australia and New Zealand. 
*A. monocephala Boiss. & Bal. in Boiss. Turkey. 
A. multifida (DC.) Boiss. Turkey. 
A. nana L. C. Europe. 
A. nobilis L. {A. neilrichii A. Kerner). S. and C. Europe, 

European Russia, W. Siberia, Turkey, Caucasus, Iran, C. 

Asia. 
A. ochroleuca Ehrh. E. Europe. 



A. odorata L. C. and SW Europe, N. Africa in Morocco and 

Algeria. 
A. oligocephala DC. Turkey, Middle East, Iraq, Iran. 
A. oxyloba (DC.) Schultz-Bip. C. and E. Europe. 
A. oxyodonta Boiss. Iran. 
*A. pachycephala Rech. f. Iran. 
A. pannonica Scheele. C, E. and SE Europe. 
A. phrygia Boiss. & Bal. in Boiss. Turkey. 
A. pindicola Hausskn. SE Europe (Greece). 
*A. pseudoaleppica Huber-Mor. Turkey. 
A. ptarmica L. Widespread in Eurasia, introduced in N. 

America. 
A. ptarmicifolia (Willd.) Rupr. ex Heimerl. Caucasus. 
A. ptarmicoides Maxim. E. Siberia, Far East, China, Japan. 
A. pyrenaica Sibth. ex Godron in Gren. & Godron. SW 

Europe. 
A. roseo-alba Ehrend. C. Europe. 
*A. sachokiana Sosn. Caucasus. 
A. salicifolia Besser. European Russia, Siberia, C. Asia, 

China. 
A. santolina L. Throughout N. Africa, Middle East, Iraq, 

Pakistan. 
A. santolinoides Lagasca. SW Europe, N. Africa in Morocco 

and Algeria. 
*A. schischkinii Sosn. Turkey, Caucasus. 
*A. sedelmeyeriana Sosn. Caucasus. 
*A. serbica Nyman {A. schurii Schultz-Bip.). SE Europe. 
A. setacea Waldst. & Kit. S., C. and SE Europe, European 

Russia, S. Siberia, Turkey, Iran, Afghanistan, C. Asia, 

China. 
A. sibirica Ledeb. Siberia, Japan, N. America in Alaska and 

Canada. 
A. sieheana Stapf. Turkey. 
A. sintenisii Huber-Mor. Turkey. 
A. sipikorensis Hausskn. & Bornm. Turkey. 
*A. spinulifolia Fenzl ex Boiss. Turkey. 
A. stricta (Koch) Schleicher ex Gremli. C. Europe. 
A. talagonica Boiss. (^4. oxylepis Boiss. & Hausskn. in 

Boiss.). Iran. 
A. tanacetifolia All. Europe. 

A. taygetea Boiss. & Heldr. in Boiss. SE Europe (Greece). 
A. tenuifolia Lam. Turkey, Caucasus, Iran. 
*A. teretifolia Willd. Turkey. 
A. thracica Velen. E. Europe. 
A. tomentosa L. SW Europe. 
A. umbellata Sibth. &. Smith. SE Europe (Greece). 
A. vermicularis Trin. Turkey, Caucasus, Iraq, Iran. 
*A. virescens (Fenzl) Heimerl in A. Kerner. SC Europe. 
A. wilhelmsii Koch {A. kermanica Gand.). Turkey, Cauca- 
sus, Syria, Iraq, Iran, Afghanistan, Pakistan, C. Asia. 
*A. wilsoniana Heimerl ex Hand.-Mazz. China. 

5L ANACYCLUS L., Sp. pL: 892 (1753). Type 
species: A. valentinus L. 

Annual or perennial herbs. Leaves alternate, rarely rosulate, 
pinnatisect. Capitula solitary or laxly corymbose, peduncu- 
late, rarely closely aggregated, radiate or discoid. Receptacle 
flat to conical, paleate. Ray florets female, fertile; tube 
flattened, persistent on the cypselas; limb white or yellow, 
abaxially sometimes reddish. Disc corolla 5-lobed, sometimes 
slightly zygomorphic with 2 larger lobes; tube somewhat 
flattened and adaxially slightly saccate. Cypselas dorsiven- 
trally flattened and laterally winged, rather thick-walled. 



130 

apically sometimes coroniform, sometimes with myxogenic 
cells; wings thick and sclerenchymatic. True pappus absent. 
Amides present. 

Distribution. Mainly W. Mediterranean; N. Africa, S. 
Europe and the Middle East. - 12 spp. 

Anacyclus was revised and discussed in detail by Humphries 
(1979), who also indicated Leucocyclus as the sister group. 

A. davatus (Desf.) Pers. 

A. homogamos (Maire) Humphries 

A. inconstans Pomel 

A. latealatus Huber-Mor. 

A. linearilobus Boiss. & Reuter 

A. maroccanus (Ball) Ball 

A. monanthos (L.) Thell. {A. cyrtolepidiodes Pomel) 

A. nigellifolius Boiss. 

A. officinarum Hayne 

A. pyrethrum (L.) Lagasca 

A. radiatus Lois. 

A. valentinus L. 

52. LEUCOCYCLUS Boiss. in Diagn. pi. orient. I 
(II): 14 (1849). Type species: L. formosus Boiss. 

A perennial herb. Leaves alternate, vermiform, pinnatisect. 
Capitula solitary, pedunculate, radiate. Receptacle flat to 
convex, paleate. Ray florets female, fertile; tube flattened, 
both adaxially and abaxially vaginate around top of cypsela; 
limb white. Disc corolla 5-lobed; tube flattened, both adaxi- 
ally and abaxially but not laterally vaginate around top of 
cypsela. Cypselas dorsiventrally flattened and laterally 
winged, rather thick-walled; wings thick and sclerenchymatic. 
Pappus absent. 

Distribution. SW Asia in Turkey. - Monotypic. 

This monotypic genus is the sister group of Anacyclus, as 
shown by Humphries (1979). Recent work by Valent- 
Vetschera (1982) has indicated that Leucocyclus is similar in 
flavonoid chemistry to certain members of Achillea sect. 
Santolinoidea. This is opposed to cypsela morphology, group- 
ing Leucocyclus with Anacyclus, and requires further investi- 
gation. 

53. MECOMISCHUS Cosson ex Benth. in Benth. & 
Hook, f.. Gen. pi. 2: 418 (1873). Type species: M. 
geslini (Cosson) Cosson (M. pedunculatus (Cosson 
& Durieu) Maire). 

Annual or perennial herbs. Leaves alternate or sometimes 
partly opposite, few-lobed or entire. Capitula solitary, 
pedunculate, radiate. Receptacle paleate; paleae with a cen- 
tral resin canal. Ray florets neuter; limb white or yellow. Disc 
corolla 5-lobed, adaxially slightly saccate, with a rather long 
tube and a partly enervate limb. Cypselas with 1 adaxial and 2 
lateral vascular strands, thin-walled, with myxogenic cells. 
Pappus absent. 

Distribution. N. Africa in Morocco and Algeria. - 2 spp. 

Mecomischus with two rather different species appears to be 
related to Chamaemelum, Cladanthus, and Rhetinolepis . 
They all have thin-walled, obovoid, myxogenic cypselas. 

M. halimifolius (Munby) Hochr. 



K. BREMER AND C. J. HUMPHRIES 
M. pedunculatus (Cosson & Durieu) Maire 

54. CHAMAEMELUM Miller in Card. Diet. abr. 4th 
edn. (1754). Lectotype: C. nobile (L.) AH. - 
Ormenis (Cass.) Cass. 

Annual or perennial herbs or half-shrubs. Leaves alternate, 
pinnatifid or variously pinnatisect. Capitula solitary or laxly 
corymbose, pedunculate, radiate, disciform, or discoid. 
Receptacle conical or elongated, paleate; paleae flat or often 
canaliculate, sometimes enclosing florets, often with a central 
resin canal, glabrous or abaxially pilose. Ray florets female, 
fertile or sterile; limb white or yellow. Disc corolla 5-lobed, 
basally saccate around the cypsela especially adaxially, with a 
rather long tube and a more or less distinct, generally 
enervate limb. Cypselas obovoid, with 1 adaxial and 2 lateral 
very thin ribs with vascular strands, thin-walled and covered 
with myxogenic cells in longitudinal rows. Pappus absent. 
Amides and particular thiophene derivatives present. 

Distribution. Mediterranean, from the Canaries in N. 
Africa and S. Europe to the Middle East. - 6 spp. 

The species of Chamaemelum were formerly placed in Orme- 
nis or as species oi Anthemis s. 1. The generic name Chamae- 
melum of Miller is prior to Cassini's Ormenis, however. 
Cassini distinguished Ormenis from Anthemis by the basally 
saccate corolla. Later, it has also been shown that Ormenis 
has a specialized cypsela morphology, different from that of 
Anthemis (Briquet, 1916). C. fuscatum, C. mixtum, and C. 
nobile (including Ormenis santolinoides (Munby) Harling) 
also differ in embryology; in contrast to Anthemis they have 
the normal monosporic type of embryo sac development 
(Harling, 1960). When Miller described the genus he 
included several species now in other genera. Only two of his 
species are presently classified in Chamaemelum, C. nobile 
and C. mixtum. The former is more well-known and the latter 
is the type of Ormenis, hence our choice of C. nobile as type 
species of Chamaemelum. In Flora europaea (Tutin et al., 
1976) the three widespread species are recognized under 
Chamaemelum (C. fuscatum, C. mixtum, C. nobile). There 
are also a number of North African species described from 
Morocco. Those that seem distinct are transferred from 
Ormenis to Chamaemelum following recent treatments of 
Benedi Gonzalez (1986, 1988^, ft; see list of species). The 
North African species are still in need of revision. 

C. eriolepis (Cosson ex Maire) Benedi. 

*C. flahaulti (Emb.) Benedi. 

C. fuscatum (Brot.) Vase. {Anthemis fuscata Brot., Anthemis 

praecox Link, Ormenis praecox (Link) Briq. & Cavill.) 
C. mixtum (L.) All. {Anthemis mixta L., Ormenis mixta (L.) 

Dumort.) 
C. nobile (L.) All. {Anthemis nobilis L., Ormenis nobilis (L.) 

Gay) 
C. scariosum (Ball) Benedi. 

55. RHETINOLEPIS Cosson in Bull. Soc. hot. Fr. 3: 
707 (1856). Type species: R. lonadioides Cosson. 

An annual herb, branched from the base; hairs dolabriform. 
Leaves alternate, entire or few-lobed. Capitula solitary or 
few closely together, almost sessile, comparatively small, 
discoid. Receptacle paleate; paleae scarious with a central 
resin canal, abaxially pilose. Corolla 5-lobed, adaxially shal- 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



131 



lowly saccate, with a narrow tube and a distinct, enervate 
limb. Cypselas narrowly obovoid, without ribs, with 1 adaxial 
and 2 lateral vascular strands, thin-walled, covered with 
myxogenic cells in longitudinal rows. Pappus absent. 

Distribution. N. Africa in Algeria, Tunisia, and Libya. - 
Monotypic. 

Rhetinolepis is a curious small annual very different in habit 
from its relatives Chamaemelum, Cladanthus, and Mecomis- 
chus. However, they have a similar specialized cypsela mor- 
phology, and many floral characters in common. In North 
African floras Rhetinolepis lonadioides is also often called 
Ormenis lonadioides (Cosson) Maire. 

56. CLADANTHUS Cass, in Bull. Sci. Soc. philom. 
Paris 1816: 199 (1816). Type species: C arabicus 
(L.)Cass. 

An annual herb with branches in whorls below the first 
capitulum. Leaves alternate, below the capitula whorled, 
pinnatisect. Capitula solitary, sessile, radiate. Receptacle 
conical, paleate and pilose; paleae canaliculate and half- 
enclosing cypsela and basal part of corolla, with a central 
resin canal, pilose on both sides. Ray florets female, sterile; 
limb yellow, with comparatively large apical lobes. Disc 
corolla 5-lobed, basally saccate around the cypsela especially 
adaxially; lobes with acute appendices. Cypselas obovate, 
laterally flattened with 1 lateral and 2 marginal very thin ribs 
with vascular strands, thin-walled and covered with myxo- 
genic cells in longitudinal rows. Pappus absent. Amides and 
particular thiophene derivatives present. 

Distribution. Mediterranean, S. Spain and N. Africa in 
Morocco, Algeria, Tunisia, and Libya. - Monotypic. 

This species is known as a relative of Chamaemelum, which 
has relatively plesiomorphic characters when compared to 
Cladanthus. The sister group, however, appears to be Rhetin- 
olepis, with small discoid capitula. Thus although the two 
genera seem rather different they both have a similar branch- 
ing habit and sessile capitula. 

8. ANTHEMIDINAE Dumort. emend. Bremer 
& Humphries, emend, nov. (Dumortier, Fl. 
belg.: 69 (1827) ('Anthemideae')). Type species: 
Anthemis maritima L. 

Herbae annuae vel perennes vel suffrutices. Capitula solitaria 
vel laxe corymbosa. Receptaculum paleaceum vel interdum 
epaleaceum. Corolla flosculi disci 5- vel raro 4-lobata tubo 
plerumque basaliter incrassato. Cypselae plerumque turbina- 
tae parietibus crassis. Pappus coroniformis vel auriculiformis 
vel nullus. Sacculus embryonis tetrasporus. 

Annual or perennial herbs or suffrutices. Leaves pinnatisect 
to variously lobed, rarely entire. Capitula solitary or laxly 
corymbose, radiate or rarely disciform or discoid. Receptacle 
convex to narrowly conical, paleate or sometimes epaleate. 
Ray floret limb white or yellow. Disc corolla 5- or rarely 
4-lobed; tube mostly basally swollen in fruit. Cypselas gener- 
ally turbinate and thick-walled, rarely with myxogenic cells. 
Pappus a corona, an auricle, or absent. Embryo sac tet- 
rasporic. 

Distribution (Table 18). Eurasia, N. and E. Africa, some 



Table 18 General distribution of Anthemidinae, Chrysantheminae, 
and genera. x=indigenous, o=introduced. 





N. 


Eur- 


SW 


S. 


N. 


S. 


Austr. S. 




Am. 


Asia 


Asia 


Eur. 


Afr. 


Afr.N.Zeal.Am. 


Anthemidinae 


o 


X 


X 


X 


X 


o 




Anthemis 


o 


X 


X 


X 


X 





O 


Nananthea 








X 








Chrysantheminae 




X 


X 


X 


X 


o 


o o 


Chrysanthemum 




X 


X 


X 


X 


o 


o 


Heteranthemis 








X 


X 






Ismelia 










X 






Argyranthemum 










X 







Anthemis species widespread as weeds also in the S. hemi- 
sphere. - 2 genera, 213 spp. 

Traditionally subtribe Anthemidinae represents all 
Anthemideae with a paleate receptacle. In our classification 
it is essentially restricted to the large genus Anthemis, the 
immediate relatives of which are unknown. Chrysantheminae 
(s. s.), also with thick-walled cypselas, is a possible sister 
group candidate. We have also provisionally included the 
isolated monotypic Nananthea in Anthemidinae. The reasons 
are given below in the discussion of Nananthea. 

The cladogram also includes the Chrysantheminae. If the 
matrices of both subtribes are analysed together, Nananthea 
appears as the sister group to Chrysantheminae rather than to 
Anthemis. Nananthea and the Chrysantheminae share the 
annual habit and absence of pappus. We would consider the 
tetrasporic embryo sac uniting Nananthea and Anthemis a 
stronger character than these two characters together, how- 
ever. The possible relationship of Nananthea to Anthemis is 
further discussed under the former genus. 

Clades and characters - Fig. 9, Tables 2, 19. 



Anl^ 57 Anthemis 
ii= 58 Nananthea 



li=Chl 



^ 59 Chrysanthemum 

li=Ch2=;7= 60 Heteranth 



60 Heteranthemis 
j[= 61 Ismelia 
^ 62 Argyranthemum 

Fig. 9 Cladogram of the Anthemidinae and the Chrysantheminae 
produced by the ie option in Hennig86. Cladogram length = 23, 
consistency index = 86, retention index = 76. 

Table 19 Data matrix for the Anthemidinae and Chrysantheminae. 1 
= presence, = absence, ? = missing data or not appHcable, p = 
polymorphic but scored as the plesiomorphic condition, a = 
polymorphic but scored as the apomorphic condition. 

nil 11 1 1 1 11 11 1 11 1 
145057655 71480 774271212857 7 15763461321 
532718112 51528122105610225726 30065414616 



51. Anthemis alaalllal 

58.Nananthea alaal?100 

59.Chrysanthemumal0al'?lal 
60. Heteranthemis al001?lal 
ei.Ismelia lla01?lll 

62.Argyranthemumlla0l?l01 



laalapppOOOOOOOOOOOO 
170001 a 1000000000000 
00000101111000000000 
10000101110111110000 
00000101110110001100 
000000011 101 lOOOOpll 



?OOpppppOOO 
????0000p00 
????00000p0 
????0000000 
????0000000 
????00000pp 



Clade Anl - subtribe Anthemidinae 
175 Embryo sac tetrasporic. 



132 



K. BREMER AND C. J. HUMPHRIES 



Anthemis 

11 Plants with dolabriform hairs. Many species oi Anthemis 
have T- or Y-shaped hairs (Napp-Zinn & Eble, 1980), which 
occur also in Artemisiinae, Leucantheminae and some genera 
of Achilleinae. 

45 Receptacle paleate. Some species of Anthemis, subgenus 
Ammanthus, have epaleate receptacles. 

82 Disc corolla tube thickened in fruit. A thickened corolla 
tube is characteristic also of many genera in Achilleinae, 
Leucantheminae and Matricariinae. 

108 Cypselas turbinate. 

Nananthea 

1 Plants annual. 

51 reversed. Floral parts without resin canals. 

72 Disc corolla 4-lobed. This character occurs also within 
Thaminophyllinae and Matricariinae. 

152 reversed. Cypsela wall not several cell layers thick, not 
partially or completely sclerified. 

172 Pappus absent in ray and disc cypselas. 

57. ANTHEMIS L., Sp. pL: 893 (1753). Type species: 
A. maritima L. 

Annual or perennial herbs or half-shrubs; indumentum fre- 
quently of dolabriform hairs. Leaves alternate, pinnatisect to 
variously lobed, occasionally entire. Capitula solitary or laxly 
corymbose, pedunculate, radiate or discoid. Receptacle con- 
vex to narrowly conical, paleate, rarely basally or totally 
epaleate; paleae scarious and oblong or subulate, truncate to 
acute to acuminate, occasionally with a central resin canal. 
Ray florets female, fertile, or neuter; limb white or rarely 
yellow or reddish. Disc corolla 5-lobed, yellow or rarely 
reddish; tube basally much swollen in fruit, rarely pilose; 
lobes rarely with acute appendices. Cypselas generally turbi- 
nate, smooth to prismatic to c. 10-ribbed, sometimes dor- 
siventrally compressed, sometimes tuberculate, thick-walled, 
rarely with myxogenic cells. Pappus a shallow, often adaxially 
more developed corona, or an adaxial auricle, or absent. 
Embryo sac tetrasporic. 

Distribution: Europe, Asia, and N. Africa, mainly in S. 
Europe and SW Asia, one species also in tropical E. Africa 
{A. tigrensis) and 2-3 species widespread as weeds also in N. 
America and the S. hemisphere {A. arvensis, A. cotula, A. 
tinctoria). - 211 spp. 

The large genus Anthemis is, despite its size, morphologically 
homogeneous and the numerous species stick together in a 
number of groups. As a whole the genus also appears to be 
monophyletic based on its turbinate (obconical, in some 
derived species more obovoid), thick-walled fruits and the 
invariably basally swollen corolla tube. The south-east Euro- 
pean species with an epaleate receptacle, hence earlier classi- 
fied in a separate genus Ammanthus, have now been 
transferred to Anthemis (Greuter, 1968). Ammanthus is 
treated by Fernandes in Flora europaea (Tutin et al., 1976) as 
a subgenus and it represents a monophyletic group of derived 
species within Anthemis. The infrageneric classification of 
Anthemis follows the common pattern with recognition of a 



number of apomorphic sections and subgenera in addition to 
a plesiomorphic and probably paraphyletic subgenus Anthe- 
mis. Yavin (1970, 1972) has proposed an elaborate infrage- 
neric classification of Anthemis. She also revised section 
Maruta. 

Anthemis was typified with A. maritima by Britton & 
Brown (1913). Later Green (in Hitchcock & Green, 1929) 
proposed A. arvensis as type species, since it is a well-known 
species also occurring in Sweden, the home-country of Lin- 
naeus. Both species fit Linnaeus' generic description in 
Genera plantarum (Linnaeus, 1757). Linnaeus did not coin 
the generic name; it was adopted from MicheH (1729) who 
included A. maritima but not A. arvensis. Hence, we consider 
the original typification with A. maritima by Britton & Brown 
to be the correct choice. 

A. aaronsohnii Eig. Middle East. 

*A. abagensis Fed. Caucasus. 

A. abrotanifolia (Willd.) Guss. Greece. 

A. aciphylla Boiss. Turkey. 

*A. adonidifolia Boiss. Turkey. 

*A. aeolica Lojac. Italy. 

A. aetnensis Schouw in Sprengel. Italy. 

A. alpestris (Hoffsgg & Link) R. Fernandes. SW Europe. 

A. altissima L. S. Europe, Krym, Turkey, Caucasus, Iraq, 

Iran, Afghanistan, C. Asia. 
A. amblyolepis Eig. Turkey, Cyprus, Middle East. 
A. ammanthus Greuter. Greece. 
A. ammophila Boiss. & Heldr. in Boiss. Turkey. 
A. anatolica Boiss. Greece, Turkey. 
A. anthemiformis (Freyn & Sint.) Grierson. Turkey. 
*A. antilibanotica Eig. Middle East. 
A. antitaurica Grierson. Turkey. 
A. arenicola Boiss. Turkey. 

*A. argyrophylla (Hal. & Georgiev) Velen. Bulgaria. 
A. armeniaca Freyn & Sint. Turkey. 
A. arvensis L. Europe, N. Africa, W. Asia, also widespread 

as a weed in N. America, S. Africa, Australia and New 

Zealand. 
*A. atropatana Iranshar. Iran. 
A. auriculata Boiss. SE Europe, Turkey. 
A. austriaca Jacq. C. and E. Europe, Krym, Turkey, Cauca- 
sus, Iran. 
A. austro-iranica Rech. f., Aellen & Esfand. Iran. 
A. bornmuelleri Stoy. & Acht. (.4. galilaea Eig). Egypt, 

Middle East. 
*A. bourgaei Boiss. & Reuter. Spain, Morocco. 
A. boveana Gay. N. Africa in Morocco, Algeria and Libya. 
*A. brachmannii Boiss. & Heldr. in Boiss. Greece. 
A. brachycarpa Eig. Middle East. 
A. br achy Stephana Bornm. & Gauba. Iran. 
A. brevicuspis Bornm. {A. feinbruniae Eig., A. rayatensis 

Eig). Middle East, Iraq, Iran. 
*A. breviradiata Eig. Middle East. 
*A. bulgarica N. N. Thin. Bulgaria. 
*A. bushehrica Iranshahr. Iran. 
A. calcarea Sosn. Turkey, Caucasus. 

A. candidissima Willd. ex Sprengel. Caucasus, Iran, C. Asia. 
A. carpatica Waldst. & Kit. ex Willd. C. and S. Europe. 
A. chia L. SE Europe, Middle East, Egypt. 
A. chrysantha Gay. Spain, Algeria. 
A. coelopoda Boiss. SE Europe, Turkey, Middle East, Iran, 

Afghanistan. 
A. cornucopiae Boiss. Middle East. 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



133 



A. corymbulosa Boiss. & Hausskn. in Boiss. Middle East. 

A. cotula L. Europe, N. Africa, W. Asia, also widespread as 
a weed in America, S. Africa, Australia and New Zealand. 

*A. cretacea Zefirov. Krym, Caucasus. 

A. cretica L. {A. anahytae Woronow ex Sosn., A. iberica M. 
Bieb., A. montana L., A. panachaica Hal., A. pindicola 
Heldr. ex Hal., A. ptarmiciformis K. Koch, A. tempskyana 
Freyn & Sint.). S. Europe, N. Africa, Turkey, Middle 
East, Iran. 

A. cuneata Huber-Mor. & Reese. Turkey. 

*A. cypria Boiss. Cyprus. 

A. cyrenaica Cosson. Libya. 

A. damascena Boiss. & Gaill. Middle East. 

A. davisii Yavin. Turkey. 

A. debilifolia Eig. Middle East. 

A. deserticola H. Kraschen. & Popov. C. Asia. 

A. deserti-syriaci Eig. Middle East. 

*A. didymaea Mout. Middle East. 

A. dipsacea Bornm. Turkey. 

*A. dubia Steven. Krym. 

A. edumea Eig. Middle East. 

A. eliezrae Eig. Middle East, Egypt. 

A. emasensis Eig. Middle East. 

A. emiliae Sosn. Caucasus. 

A. filicauUs (Boiss. & Heldr.) Greuter. Greece. 

A. fimbriata Boiss. Turkey. 

A. flexicaulis Rech. f. Greece. 

A. freitagii Iranshahr. Afghanistan. 

A. fruticulosa M. Bieb. Caucasus. 

A. fulvida Grierson. Turkey. 

A. fumariifolia Boiss. Turkey. 

*A. fumarioides Hochst. Middle East. 

M. fungosa Boiss. & Hausskn. Iran. 

A. gaudium-solis Velen. Bulgaria. 

A. gay ana Boiss. Iran. 

A. gerardiana Jordan. France. 

A. gilanica Boiss. Iran. 

*A. gilletti Iranshahr. Iraq, Iran. 

A. glaberrima (Rech. f.). Greuter. Greece. 

A. glareosa Durieu & Barratte. Libya. 

*A. gracilis Iranshahr. Iran. 

A. gross heimii Sosn. Caucasus. 

A. halophila Boiss. & Bal. in Boiss. Turkey. 

*A. hamrinensis Iranshahr. Iraq. 

A. handel-mazzettii Eig. Middle East. 

A. haussknechtii Boiss. & Reuter in Boiss. Syria, Iraq, Iran. 

A. hebronica Boiss. & Kotschy. Middle East, Egypt. 

A. hemistephana Boiss. Middle East, Iran. 

*A. hermonis Eig. Middle East. 

*A. hinkovae N. N. Thin. Bulgaria. 

A. hirtella Winkler. C. Asia. 

A. homalolepis Eig. Middle East. 

A. hyalina DC. Turkey, Middle East, Iraq, Iran. 

A. hydruntina Groves. Italy. 

A. indurata Del. Middle East, N. Africa in Egypt and Libya. 

A. ismelia Lojac. Italy. 

*A. jailensis Zefirov. Krym. 

M. jordanovii Stoy. & Acht. Bulgaria. 

A. kandaharica Iranshahr. Afghanistan, Pakistan. 

*A. karabaghensis Mikheev. Caucasus. 

A. kitanovii N. N. Thin. Bulgaria. 

A. kitenensis N. N. Thin. Bulgaria. 

A. kotschyana Boiss. Turkey, Middle East, Iraq, Iran. 

*A. krugeriana Pampan. Libya. 



*A. kurdica Iranshahr. Iraq. 

*A. kuzmanovii N . N. Thin. Bulgaria. 

A. laconica R. Franzen. Greece. 

*A. leptophylla Eig. Iraq, Iran. 

A. leucanthemifolia Boiss. & Blanchet. Middle East, Egypt. 

A. leucolepis Eig. Middle East. 

M. linczevskyi Fed. C. Asia. 

*A. lithuanica (DC.) Besser ex Trautv. Europe in Lithuania 

and Russia. 
A. lores tanica Iranshahr. Iran. 

A. lyonnetioides (Boiss. & Kotschy) Boiss. Middle East. 
*A. macedonica Boiss. & Orph. in Boiss. SE Europe. 
A. macrantha Heuffel. SE Europe. 
*A. macroglossa Sommier & Levier. Caucasus. 
A. maris-mortui Eig. Middle East. 
M. maris-nigri Fed. Caucasus. 
A. maritima L. SW Europe, N. Africa. 
*A. markhotensis Fed. Caucasus. 
A. marschalliana Willd. Caucasus. 
A. mauritiana Maire & Sennen. Morocco. 
A. mazandaranica Iranshahr. Iran. 

A. melampodina Del. {A. deserti Boiss.) Middle East, Egypt. 
A. melanacme Boiss. & Hausskn. in Boiss. Turkey, Middle 

East. 
A. melanoloma Trautv. Turkey. 
*A. meteorica Hausskn. Greece. 
A. micrantha Boiss. & Hausskn. Iraq. 
A. microcephala (Schrenk) B. Fedtsch. {A. straussii Bornm., 

A. tenuiflora Gilli). C. Asia, Iraq, Iran, Afghanistan. 
A. microlepis Eig. Middle East. 

A. microsperma Boiss. & Kotschy. Middle East, Egypt. 
M. mirheydari Iranshahr. Iran. 
A. moghanica Iranshahr. Iran. 
M. monantha Willd. Krym. 
*A. monilicosta Pomel. N. Africa in Morocco, Algeria and 

Libya. 
A. muricata (DC.) Guss. Italy. 
*A. nabataea Eig. Middle East. 
A. odontostephana Boiss. {A. tubicina Boiss. & Hausskn. in 

Boiss.). Iran, C. Asia. 
*A. orbelica Pancic. Bulgaria. 
A. orientalis (L.) Degen {A. pectinata (Bory & Chaub.) 

Boiss. & Reuter). Greece, Turkey. 
A. oxylepis (Boiss.) Boiss. Turkey. 
A. palestina Reuter in Boiss. {A. melanolepis Boiss., A. 

syriaca Bornm.). Turkey, Cyprus, Middle East, SE 

Europe. 
A. parnassica (Boiss. & Heldr.) R. Fernandes. SE Europe. 
A. parnesia Boiss. & Heldr. in Boiss. Greece. 
*A. parviceps Dobrocz. & Fed. Krym. 
A. parvifolia Eig. Middle East. 
*A. patentissima Eig. Middle East. 
A. pauciloba Boiss. Turkey, Iraq. 
A. pedunculata Desf. N. Africa in Morocco, Algeria and 

Libya. 
A. persepolitana Boiss. Iraq, Lebanon, Syria. 
A. persica Boiss. Iran. 
A. pestalozzae Boiss. Turkey. 
A. plebeia Boiss. & Noe. Iraq. 
M. plutonia Meikle. Cyprus. 
A. pseudocotula Boiss. {A. behboudiana Rech. f. & Esfand.). 

Turkey, Cyprus, Middle East, Iraq, Iran, N. Africa in 

Egypt and Libya. 



134 



K. BREMER AND C. J. HUMPHRIES 



A. punctata Vahl. Italy, N. Africa in Morocco, Algeria and 

Tunisia. 
*A. pungens Yavin. Turkey. 
A. rascheyana Boiss. Middle East. 
*A. regis-borisii Stoy. & Acht. Bulgaria. 
A. retusa Del. Egypt. 
*A. rhodensis Boiss. Turkey. 

*A. rhodocentra Iranshahr. Iran, Afghanistan, Pakistan. 
A. rigida (Sibth. & Smith) Boiss. & Heldr. SE Europe, 

Turkey, Cyprus. 
A. rosea Smith in Sibth. & Smith. Turkey. 
A. rumelica (Velen.) Stoy. & Acht. Bulgaria. 
A. ruthenica M. Bieb. C. and SE Europe, Caucasus. 
A. sabuUfolia Pomel. N. Africa. 
A. saguramica Sosn. Caucasus. 
A. samuelssonii Rech. f. Middle East. 
A. sancti-johannis Turrill. Bulgaria. 
*A. saportana Albov. Caucasus. 
A. scaettae Pampan. Libya. 
A. scariosa Banks & Sol. in Russell. Turkey, Middle East, 

Iraq, Iran. 
*A. schischkiniana Fed. Caucasus. 
A. schizostephana Boiss. & Hausskn. Iraq, Iran. 
A. scopulorum Rech. f. Greece. 
A. scrobicularis Yavin. Middle East. 
A. secundiramea Biv. SW Europe, N. Africa in Algeria and 

Tunisia. 
A. segetalis Ten. {A. brachycentros Gay ex W. Koch). S. 

Europe. 
A. semiensis Pichi-Serm. Ethiopia. 
A. sibthorpii Griseb. Greece. 
A. sintenisii Freyn. Turkey. 
A. sosnovskyana Fed. Caucasus. 
A. spruneri Boiss. & Heldr. in Boiss. Greece. 
A. sterilis Steven. Krym. 
A. stiparum Pomel. N. Africa. 
*A. stribrnyi Velen. Bulgaria. 
M. susiana Nab. Iraq, Iran. 
A. talyschensis Fed. Caucasus, Iran. 
*A. taubertii Durieu & Barratte. Libya. 
A. tenuicarpa Eig. Middle East. 
A. tenuiloba (DC.) R. Fernandes. SE Europe. 
A. tigrensis Gay ex A. Richards. E. Africa. 
A. tinctoria L. (y4. debilis Fed., A. euxina Boiss., A. subtinc- 

toria Dobrocz.). Europe, W. and C. Asia, also naturalized 

in N. America. 
A. tomentella Greuter. Greece. 

A. tomentosa L. {A. peregrina L.). SE Europe, Turkey. 
*A. tranzcheliana Fed. Krym. 
A. tricolor Boiss. Cyprus. 
A. tricornis Eig. Turkey. 

A. tripolitana Boiss. & Blanchet in Boiss. Middle East. 
A. triumfettii (L.) DC. in Lam. & DC. (A. dumetorum Sosn., 

A. khorassanica Rech. f., ^4. rigescens Willd.). S. Europe, 

Turkey, Caucasus, Iran. 
A. trotzkiana Claus ex Bunge. S. Russia, C. Asia. 
A. tuberculata Boiss. Spain. 
*A. virescens Velen. Bulgaria. 
A. wallii Huber-Mor. & Reese. Turkey. 
*A. werneri Stoy. & Acht. Greece. 
A. wettsteiniana Hand.-Mazz. Iraq, Iran. 
A. wiedemanniana Fischer & C. Meyer. Caucasus, Turkey. 
A. woronowii Sosn. Caucasus. 
A. xylopoda O. Schwarz. Turkey. 



A. yemenensis Podl. Yemen. 

M. zephyrovii Dobrocz. Krym. 

A. zoharyana Eig. Middle East, Egypt. 

A. zyghia Woronow. Caucasus. 

58. NANANTHEA DC, Prodr. 6: 45 (1838). Type 
species: N. perpusilla (Lois.) DC. 

A delicate, somewhat succulent, annual herb. Leaves alter- 
nate, pinnatifid, with obovate lobes. Capitula solitary, long- 
pedunculate, very small (2-5 mm diam.), radiate or 
disciform. Involucre of 5-10 wide bracts in 1 to 2 rows. 
Receptacle conical, epaleate. Ray florets female, fertile, with 
or without a white lamina. Disc corolla 4-lobed, with a short 
tube and large lobes. Cypselas obovoid, with myxogenic cells; 
ray cypselas with 2 lateral vascular strands; disc cypselas with 
2 lateral, 1 adaxial and 1 pseudolateral vascular strand. 
Pappus absent. Embryo sac tetrasporic. 

Distribution. S. Europe, Corsica and Sardinia. - Mono- 
typic. 

Nananthea is a genus of uncertain position. Traditionally it 
has been included in a widely circumscribed and heteroge- 
neous Cotula group but it shares no synapomorphies with the 
Cotula group, which is part of subtribe Matricariinae, as 
understood here. Nananthea is here provisionally placed 
together with Anthemis, following a suggestion by Reitbrecht 
(1974). It could be a highly speciaHzed Anthemis derivative, 
related to part of Anthemis, possibly subgenus Ammanthus. 
Both genera have tetrasporic embryo sacs. This also occurs in 
Tanacetum, Tripleurospermum, and Heteranthemis, but Nan- 
anthea is hardly reminiscent of these genera. 

9. CHRYSANTHEMINAE Less, emend. Bremer 
& Humphries, emend, nov. (Lessing in Linnaea 
6: 167 (1831) ('Chrysanthemeae')). Type species: 
Chrysanthemum coronarium L. 

Herbae annuae vel perennes vel suffrutices vel frutices. 
Capitula solitaria vel laxe corymbosa. Bracteae involucri 
latae, plurinerves. Receptaculum epaleaceum. Cypselae pari- 
etibus crassis, heteromorphae; cypselae flosculorum radii 
triquetrae, alatae; eae flosculorum disci plerumque lateraliter 
compressae et abaxialiter adaxialiterque alatae vel raro ter- 
etes ad prismaticae. Pappus nullus. 

Annual or perennial herbs, half-shrubs or shrubs. Leaves 
serrate-dentate-pinnatifid to variously dissected. Capitula 
solitary or laxly corymbose, pedunculate, radiate. Involucral 
bracts wide, many- veined. Receptacle convex to conical, 
epaleate. Ray floret limb white and/or yellow. Disc corolla 
5-lobed, yellow or rarely red. Cypselas thick-walled, without 
myxogenic cells, heteromorphic; ray cypselas triquetrous, 
winged; disc cypselas generally laterally flattened and abaxi- 
ally and adaxially winged, or rarely terete to prismatic. 
Pappus absent. 

Distribution (Table 18). Eurasia, N. Africa and Macaron- 
esia. Chrysanthemum coronarium widespread as a weed also 
in the S. hemisphere. - 4 genera, 28 spp. 

Traditionally subtribe Chrysantheminae comprises all of the 
Anthemideae species with an epaleate receptacle. As circum- 
scribed here it is a small and homogeneous taxon. The close 
relationship between Chrysanthemum s. s. and the three 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 

other genera of this subtribe has also been recognized by 
earher authors and is discussed by Humphries (1976). 

Clades and characters - Fig. 9, Tables 2 and 19. 

Clade Chi - subtribe Chrysantheminae 

1 Plants annual. Argyranthemum consists of shrublets or 
half-shrubs, here considered a secondary development within 
the subtribe. A shrubby habit is commonly evolved in island 
groups. 

41 Involucral bracts wide, flabelliform. This character also 
occurs in several genera of other subtribes. 

120 Cypselas heter amorphic; ray cypselas triquetrous, winged; 
disc cypselas terete to prismatic to laterally flattened. 

172 Pappus absent in ray and disc cypselas. 

Chrysanthemum 

52 reversed. Ray floret limb not white, but yellow. 

75 Disc corolla lobes with central resin sacs. 

Clade Ch2 

107 reversed. Cypselas not terete to weakly angled, but acutely 
angled. 

116 Disc cypselas laterally flattened. 

121 Disc cypselas abaxially and adaxially winged. 
Heteranthemis 

10 Plants covered with viscid hairs. 

52 reversed. Ray floret limb not white, but yellow. 

122 Cypsela wings as apical spines. 

175 Embryo sac tetrasporic. The embryo sac of Ismelia is 
monosporic (Hading, 1951). 

182 Flavonol 5-glycosides present. Most of the related genera 
have not been investigated chemically. 

Ismelia 

55 Ray floret limb deeply emarginate. 

77 Disc corolla red. 

Argyranthemum 

1 reversed. See clade Chi. 

2 Plants shrubby. See clade Chi under character 1. 
51 reversed. Floral parts without resin canals. 

176 Embryo sac disporic. 

59. CHRYSANTHEMUM L., Sp. pi.: 887 (1753). 
Type species: C coronarium L. 

Annual herbs. Leaves alternate, deeply serrate-dentate and 
pinnatifid to pectinate, somewhat amplexicaul. Capitula soli- 
tary or laxly corymbose, pedunculate, radiate. Involucral 
bracts wide, many- veined, with resin canals. Receptacle 
convex, epaleate. Ray florets female, fertile; limb yellow or 
white distally, many-veined. Ray cypselas triquetrous, later- 
ally winged, adaxially with a narrow wing or ribbed; pappus 
absent. Disc corolla 5-lobed; lobes with central resin sacs. 
Disc cypselas prismatic with a narrow adaxial wing or terete. 



135 

with a thick undulating wall, thus apparently ribbed; pappus 
absent. 

Distribution. Europe, Asia and N. Africa, C. coronarium 
widespread as a weed. - 2 spp. 

The adaxial cypsela wings of C. coronarium are similar to 
those in Heteranthemis, Ismelia, and Argyranthemum, 
whereas C. segetum has terete disc cypselas and only laterally 
winged ray cypselas. The adaxial cypsela wing is conceived as 
a parallelism, since the two species of Chrysanthemum are 
united for example by their corolla lobe resin sacs. 

C. carinatum Schousboe is more closely related to Heteran- 
themis and Argyranthemum than to the two Chrysanthemum 
species adopted here, and thus it is transferred to Ismelia. 

C. coronarium L. 
C. segetum L. 

60. HETERANTHEMIS Schott in Isis, Oken 1818 (5): 
822 (1816). Type species: H. viscidehirta Schott. 

An annual herb covered with viscid glandular hairs. Leaves 
alternate, serrate-dentate to pinnatifid. Capitula solitary or 
laxly corymbose, pedunculate, radiate. Involucral bracts 
wide, many-veined, with resin canals. Receptacle convex, 
epaleate. Ray florets female, fertile; limb yellow, many- 
veined. Ray cypselas triquetrous, laterally and adaxially 
winged; wings projected to apical spines; pappus absent. Disc 
corolla 5-lobed. Disc cypselas laterally flattened, winged; 
adaxial wing projected to an apical spine; pappus absent. 
Embryo sac tetrasporic. 

Distribution. SW Europe in Spain and Portugal, N. Africa 
in Morocco and Algeria. - Monotypic. 

This species, also known as Chrysanthemum viscidehirtum 
(Schott) Thell., is distinguished from Ismelia and Argyranthe- 
mum by its pubescence of viscid glandular hairs and the apical 
spines on the cypsela wings. 

61. ISMELIA Cass, in Diet. ScL Nat. 41: 40 (1826). 
Type species: /. versicolor Cass. (/. carinata 
(Schousboe) Schultz-Bip.). 

An annual herb. Leaves alternate, pinnatisect. Capitula 
soHtary or laxly corymbose, pedunculate, radiate. Involucral 
bracts wide, many-veined, with resin canals. Receptacle 
convex to conical, epaleate. Ray florets female, fertile; limb 
yellow, basally reddish or white, many-veined, deeply emar- 
ginate. Ray cypselas triquetrous, laterally and adaxially 
winged; pappus absent. Disc corolla 5-lobed, red to purple. 
Disc cypselas laterally flattened, winged; pappus absent. 

Distribution. N. Africa in Morocco but frequently escaped 
from cultivation. - Monotypic. 

This handsome species, frequently cultivated as an ornamen- 
tal, is commonly known as Chrysanthemum carinatum 
Schousboe. It is, however, more closely related to Heteran- 
themis and Argyranthemum than to Chrysanthemum coro- 
narium and Chrysanthemum segetum, the two species here 
retained in that genus. Ismelia, Heteranthemis, and Argyran- 
themum form a monophyletic group based on their laterally 
compressed disc cypselas and their especially strongly devel- 
oped cypsela wings on the adaxial side. Interestingly, Schultz- 
Bipontinus (1844/?) expanded the concept of Ismelia to 



136 



K. BREMER AND C. J. HUMPHRIES 



include various Canary Island endemics of Argyranthemum 
(see Humphries, 1976, for details). Argyranthemum, how- 
ever, forms a monophyletic group, so Ismelia is related to 
Argyranthemum as a whole rather than to part of it. Uniting 
Ismelia and then by consequence also Heteranthemis with 
Argyranthemum because of their similar fruits is hardly 
desirable, since it necessitates recombination of all Argyran- 
themum specific names, Argyranthemum being the youngest 
name. 

62. ARGYRANTHEMUM Webb ex Schultz-Bip. in 
Webb & Berthelot, Hist. nat. lies Canaries 3 (2,2): 
245, 258 (1844ft). Type species: A. frutescens (L.) 
Schultz-Bip. 

Shrublets or half-shrubs. Leaves alternate, variously dis- 
sected. Capitula solitary or laxly corymbose, pedunculate, 
radiate. Receptacle convex to conical, epaleate. Ray florets 
female, fertile; limb white, rarely yellow or pink, many- 
veined. Ray cypselas triquetrous, generally laterally and 
adaxially strongly winged, sometimes coalesced into groups; 
wings sometimes reduced, often apically projected to a 
pappus-like corona; true pappus absent. Disc corolla 5-lobed, 
yellow; lobes rarely reddish purple. Disc cypselas generally 
laterally flattened and adaxially and abaxially winged, some- 
times prismatic to terete and wingless, sometimes coalesced 
with ray cypselas, apically often coroniform; true pappus 
absent. Embryo sac bisporic. 

Distribution. Macaronesia in the Canary Islands, Madeira 
{A. dissectum, A. haematomma, A. pinnatifidum) , and the 
Salvage Islands {A. thalassophilum) . - 24 spp. 

Argyranthemum was revised by Humphries (1976). As 
already pointed out by him, it is related to Ismelia and 
Heteranthemis. Argyranthemum has, unlike these genera, 
evolved into many species with even more specialized some- 
times coalesced fruits often with folded wings or sometimes 
secondarily wingless (Borgen, 1972). A new species, A. 
sundingii, was described by Borgen (1980) and a study of 
variation within A. pinnatifidum was undertaken by Rustan 
(1981). 

A. adauctum (Link) Humphries 

A. broussonetii (Pers.) Humphries 

A. callichrysum (Svent.) Humphries 

A. coronopifolium (Willd.) Humphries 

A. dissectum (Lowe) Lowe 

A. X escarrei (Svent.) Humphries 

A. filifolium (Schultz-Bip.) Humphries 

A. foeniculaceum (Willd.) Webb ex Schultz-Bip. 

A. frutescens (L.) Schultz-Bip. 

A. gracile Schultz-Bip. 

A. haematomma (Lowe) Lowe 

A. haouarytheum Humphries & Bramwell 

A. hierrense Humphries 

A. jacobiifolium Kunkel 

A. lemsii Humphries 

A. lidii Humphries 

A. maderense (D. Don) Humphries 

A. pinnatifidum (L. f.) Lowe 

A. sundingii Borgen 

A. sventenii Humphries & Aldridge 

A. tenerifae Humphries 

A. thalassophilum (Svent.) Humphries 



A. webbii Schultz-Bip. 

A. winteri (Svent.) Humphries 

10. LEUCANTHEMINAE Bremer & 
Humphries, subtrib. nov. 

Type species: Leucanthemum vulgar e Lam. 

Herbae annuae vel perennes vel interdum suffruticosae. Folia 
saepe serrata-dentata vel interdum pectinata-lobata, trifur- 
cata vel integra. Capitula solitaria, pedunculata. Flosculi disci 
tubo plerumque incrassato. Cypselae plerumque 10(8-12) vel 
interdum pauciores costas praebentes, inter costas saepe 
lacunis vallecularibus et canalibus secretoriis et fasciculis 
vascularibus {Leucanthemum et genera sequentia), in costis 
plerumque cellulis mucilaginis, instructae. Pappus coronifor- 
mis vel adaxialiter auriculiformis vel nullus. 

Annual or perennial herbs, sometimes suffruticose; indumen- 
tum frequently of dolabriform hairs. Leaves often serrate- 
dentate, sometimes pectinate-lobed, trifurcate or entire. 
Capitula solitary, pedunculate, generally radiate or some- 
times discoid. Involucral bracts sometimes with dark brown 
margins, sometimes wide and flabelliform. Receptacle flat to 
conical, epaleate. Ray floret limb white, yellow or rarely 
reddish. Disc corolla 5-lobed or rarely 4-lobed; tube generally 
swollen in fruit. Cypselas mostly 10(8-12)-ribbed, sometimes 
with fewer ribs, often with vallecular lacunae and vallecular 
secretory canals as well as vascular strands between the ribs 
{Leucanthemum group of genera), generally with myxogenic 
cells along the ribs. Pappus a corona, an adaxial auricle, or 
absent. 

Distribution (Table 20): Eurasia and N. Africa, mainly in 
the Mediterranean region, some Leucanthemum species 
widely introduced, also in N. America and the S. hemisphere. 
- 16 genera, 75spp. 

Leucantheminae consists of one well defined monophyletic 
subclade, the Leucanthemum group of genera, a smaller 
subclade comprised of Leucanthemopsis , Hymenostemma 
and Prolongoa, as well as a number of isolated genera of 
uncertain position, here provisionally included in this sub- 
tribe. 

The Leucanthemum group of genera (71-78; see Fig. 10), 
are all characterized by their specialized cypsela wall with 
vallecular lacunae, secretory canals, and vascular strands. 
Their status as a group has been recognized by several earlier 
authors (for example, see Briquet, 1916). 

Leucanthemopsis is apparently allied to the two Iberian 
monotypic genera Hymenostemma and Prolongoa, a relation- 
ship that hitherto has not been clearly recognized. 

The subtribal description above does not cover all diver- 
gent characters of the genera, but applies mainly to the 
Leucanthemum and Leucanthemopsis groups. 

A relationship between Leucanthemopsis and Leucanthe- 
mum has been suggested several times, e. g. by Heywood 
(1954, 1976). It is supported by presence of flavonol 
5-glycosides in both genera. They are also found in Coleoste- 
phus and Plagius and constitute a possible synapomorphy for 
the Leucanthemum and Leucanthemopsis groups. Other gen- 
era of these groups have not been investigated chemically, as 
far as we know. 

Included in Leucantheminae are also a number of odd, 
mainly monotypic genera of uncertain affinity. Lepidopho- 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 

Table 20 General distribution of Leucantheminae and genera. x=indigenous, o=introduced. 



137 



N.An 


1. Eur- 


C.& 


E.SW. 


S.Eur. 


N 




Asia 


Asia 


Asia 






jucantheminae o 


X 


X 


o 


X 


X 


Lepidophorum 








X 




Nipponanthemum 




X 








Leucanthemella 


X 


X 








Nivellea 










X 


Phalacrocarpum 








X 




Leucanthemopsis 


X 






X 


X 


Hymenostemma 








X 


X 


Prolongoa 








X 




Leucanthemum o 


X 







X 


X 


Rhodanthemum 








X 


X 


Leucoglossum 








X 


X 


Chlamydophora 








X 


X 


Chrysanthoglossum 










X 


Glossopappus 








X 


X 


Coleostephus 








X 


X 


Plagius 








X 


X 



N.Afr. S.Afr. 



Austr. S.Am. 
N.Zeal. 



rum, Nipponanthemum, Leucanthemella, Nivellea and Phala- 
crocarpum are classified in this subtribe mainly because of 
their serrate-dentate leaves (not in Nivellea) but also on 
account of some floral similarities. These genera have 
10(8-12)-ribbed cypselas as in the Leucanthemum group. An 
exception is Lepidophorum with 5-ribbed cypselas. The posi- 
tions of these genera are further discussed under each genus. 
There is one most parsimonious cladogram produced 
from the data matrix. It differs from the cladogram pre- 
sented by having Hymenostemma and Prolongoa as the 
sister group to Lepidophorum rather than to Leucanthe- 
mopsis. However, we have chosen the latter arrangement, 
which is one step longer. We consider the peculiar pappus 
(character 165) shared by Hymenostemma, Prolongoa and 
Leucanthemopsis a stronger character than the annual 
habit and the loss of brown involucral bracts together 
(characters 1 and 44), which the former two genera share 
with Lepidophorum. 

Clades and characters - Fig. 10, Tables 2, 21. 

Clade Lei - subtribe Leucantheminae 



theminae, notably those provisionally included in the early 
part of the account, have dolabriform hairs. The character 
occurs also in Artemisiinae, Anthemis (Anthemidinae), and 
some genera of Achilleinae. Possibly it is a synapomorphy at 
a lower level within the tribe. 

Lepidophorum 

1 Plants annual. 

45 Receptacle paleate. 

52 reversed. Ray floret limb not white, but yellow. 

53 Ray floret limb golden yellow. Similar rays occur in 
subclade LelO of the Leucanthemum group of genera. 

133 Cypselas with costal resin canals or sacs. 

Clade Le2 

124 Cypselas with 10 (8-12) multicellular epicarpic ribs. Fewer 
ribs occur in most species of Leucanthemopsis and in 
Hymenostemma and Prolongoa. 



11 Plants with dolabriform hairs. Most genera of Leucan- Nipponanthemum 



L=Lel=n= 63 Lepidophorum 

lLLe2=7r= 64 Nipponanthemum 
[= 65 Leucathemella 
l!=Le3:i7= 66 Nivellea 



tLe4:^p= 67 Phalacrocarpum 

lL=Le5^n=Le6TJ= 68 Leucanthemopsis 

L=helTr= 69 Hymenostemma 

it= 70 Prolongoa 
rp 71 Leucanthemum 
LLesiU 72 Rhodanthemum 



=Le9 



73 Leucoglossum 

74 Chlamydophoi 

75 Chrysanthoglossum 



LeloE 74 Chlamydophora 



Jr [7= /^c^r3 

lLLeiiiL=Lel2Tf= 76 Glossopappus 
\\= 77 Coleostephus 



78 Plagius 



Fig. 10 Cladogram of the Leucantheminae produced by the bb option in Hennig86. Cladogram length = 29, consistency index = 58, 
retention index = 79. 



138 



K. BREMER AND C. J. HUMPHRIES 



Table 21 Data matrix for the Leucantheminae. 1 = presence, = absence, ? = missing data or not applicable, p = polymorphic but scored as the 
plesiomorphic condition, a = polymorphic but scored as the apomorphic condition. 



1111 111 1111111111111 1 1111 
145057652 1 4532 7 8 4 10 7 8 6 5 7 3 3 6 8 2 2 12 3 6 4 7 4 2 6 
532718111 1153342224444253714564060258131959 



111 1 

1577755556666617 
305 605 679056785 7 



63. Lepidophorum a 1 1 

64. Nipponanthemum a 1 1 1 

65. Leucanthemella a 1 1 1 

66. Nivellea 1111 

67. Phalacrocarpum a 1 1 1 

68. Leucanthetnopsis 1 1 a 1 

69. Hymenostemma 1111 

70. Prolongoa 110 1 

71. Leucanthemum a 1 a 1 

72. Rhodanthemum 1111 

73. Leucoglossum a 1 1 1 

74. Chlamydophora a 1 ? 1 

75. Chrysanthoglossum 110 1 

76. Glossopappus a 1 1 

77. Coleostephus a 1 1 

78. Plagius a 1 ? 1 



1 1 1 

1 1 

1 1 1 

1 1 

1 1 1 

1 1 

1 1 

1 1 

1 1 1 

1 1 

1 1 1 

1 1 

1 1 

I 1 1 
1 1 1 

I I 1 



? 1 1 

1 00 
1 00 
? 1 
1 00 
1 00 
1 1 
1 1 
000 
?00 
? 1 
? 1 
? aO 
1 1 
1 1 
?00 



1 1 
00 1 
00 1 
00 1 
00 1 
000 
000 
000 
00 1 
00 1 

00 1 
?0 1 

1 1 
1 1 
1 1 
?0 1 



000 
1 00 
1 
1 1 
1 1 
00 1 
00 1 
000 
Op 1 
00 1 
00 1 
00 1 
00 1 
00 1 
00 1 
00 1 



00000 
00000 
00000 
00000 
11110 
1000 1 
0000 ? 
0000 ? 
1000 1 
1000? 
0000 ? 
0000 ? 
0000 ? 
0000 ? 
0000 1 
0000 1 



000 
000 
000 
000 
000 
1 00 
1 00 

1 1 1 

000 
000 
000 
000 
000 
000 
000 
000 



000 
000 
000 
000 
000 
000 
000 
1 00 
1 1 
1 1 

1 1 1 
? 1 1 

1 1 
1 1 
1 1 

? 1 1 



000 
000 
?00 
?00 
?00 
000 
000 
000 
a 1 
1 1 
1 00 
1 00 
000 
1 00 
1 00 
1 00 



000 
000 
000 
000 
000 
000 
000 
000 
000 
1 00 
1 
1 1 
000 
000 
000 
000 



000 

000 
000 
000 
000 
000 
000 
000 
pOO 
000 
000 
1 1 
00 1 
1 

00 1 

1 00 



00000 
00000 
00000 
00000 
00000 
00000 
00000 
00000 
00000 
00000 
00000 
00000 
11000 
10 111 
1 1 aO 
10 110 



?00 

9 9 9 

9 9 9 

? ? 

9 9 9 

9 9 9 

9 9 9 

9 9 9 

? ? 

9 9 9 

9 9 9 

9 9 9 

9 9 9 

? ? 

? ? 

9 9 9 



000 
?00 
? ? 
? 
? ? 
? 00 
? 00 
? 00 
000 
? 00 
? 00 
? ? 
?00 
000 
000 
? ? 



000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
000 
? ? ? 
000 
000 
000 

9 9 9 



000 

000 
000 
000 
000 
000 
000 
000 
000 
000 
000 

9 9 9 

000 
000 
000 

9 9 9 



0000 
0000 
0000 
0000 
000 
000 
0000 
0000 
000 
OOOp 
0000 
? ? ? 
0000 
0000 
0000 

9 9 9 



2 Plants shrubby. 

51 reversed. Floral parts without resin canals. 

Leucanthemella 

172 Pappus absent in ray and disc cypselas. 

Clade Le3 

82 Disc corolla tube thickened in fruit. This character is 
sometimes vaguely expressed and absent at least in Prolon- 
goa. 

Nivellea 

1 Plants annual. 

21 reversed. Leaves not serrate-dentate. 

172 Pappus absent in ray and disc cypselas. 

Clade Le4 

44 Involucral bracts with dark brown margins. This character 
is absent in several genera (see cladogram) and present in 
many genera of other subtribes (Cancriniinae, Tanacetinae, 
Artemisiinae). It may be a synapomorphy at a lower level 
with reversals in several instances. 

Phalacrocarpum 

14 Leaves opposite. 

104 Disc floret style-branches fused. 

172 Pappus absent in ray and disc cypselas. 

\1A Testa epidermis cells thick-walled and dark reddish. 

Clade Le5 

182 Flavonol 5-glycosides present. In this subtribe only Leu- 
canthemopsis, Leucanthemum, Coleostephus and Plagius 
have been investigated for these compounds. 

Clade Le6 

21 reversed. Leaves not serrate-dentate. 

124 reversed. See clade Le2. 

165 Pappus a scarious, flimsy corona. 



Leucanthemopsis 

There is no autapomorphy for this genus. 

Clade Le7 

I Plants annual. 

44 reversed. See clade Le4. 

Hymenostemma 

There is no autapomorphy for this genus. 

Prolongoa 

52 reversed. Ray floret limb not white, but yellow. 

82 reversed See clade Le3. 

103 Disc floret style-branches long-penicillate. 

157 Cypsela wall with rod-shaped crystals in small packets. 

171 Pappus absent in disc cypselas, but present in ray cypselas. 

Clade Le8 - The Leucanthemum group of genera 

134 Cypselas with vallecular secretory canals. 

135 Cypselas with vallecular vascular strands. 

166 Pappus adaxially long. In Chrysanthoglossum the pappus 
is coroniform and hardly longer adaxially. 

Leucanthemum 

II reversed. Plants without dolabriform hairs. 
184 Anthocyanin present in root tips. 
Rhodanthemum 

20 Leaves spathulate in outline, ternate to ternately pinnate. 

21 reversed. Leaves not serrate-dentate. 

126 Cypsela ribs protruding, narrow and somewhat wing-like. 

Clade Le9 

1 Plants annual. The character reverses in Plagius, which is 
perennial. 

44 reversed. See clade Le4. 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 

Clade LelO 

110 Cypselas ellipsoid, small, c.l mm long. 

Leucoglossum 

171 Pappus absent in disc cypselas, but present in ray cypselas. 

Chlamydophora 

21 reversed. Leaves not serrate-dentate. 

22 Leaves entire or apically tridentate. 
35 Capitula discoid. 

168 Pappus a large, scarious, adaxial but basally coroniform 
auricle, as long as the corolla or longer. 

Clade Lell 

41 Involucral bracts wide, flabelliform. The character reverses 
in Plagius, where the involucral bracts are not flabelliform. 

52 reversed. Ray floret limb not white, but yellow. 

53 Ray floret limb golden yellow. 

73 Disc corolla lobes with dorsal appendages. 

Chrysanthoglossum 

21 reversed. Leaves not serrate-dentate. 

141 Cypselas with dense rows of myxogenic cells also on the 
corona. 

166 reversed. See clade Le8. 

Clade Lel2 

109 Cypselas arcuate. 

125 Cypsela ribs basally fused into a more or less well 
developed foot callus. This character is variously strongly 
expressed in the different species of the three genera 
involved. 

Glossopappus 

169 Pappus a large, scarious, adaxial, flabelliform auricle, as 
long as the corolla or longer. 

Coleostephus 

There is no autapomorphy for this genus. 

Plagius 

1 reversed. See clade Le9. 

35 Capitula discoid. 

41 reversed. See clade Lell. 

63. LEPIDOPHORUM Necker ex Cass, in Diet. Sc. 
Nat. 26: 36 (1823). Type species: L. repandum (L.) 
DC. 

An annual or biennial herb. Leaves alternate, oblong to 
obovate-spathulate, serrate. Capitula solitary, pedunculate, 
radiate. Involucral bracts wide, many-veined. Receptacle 
paleate; paleae scarious with a conspicuous resin canal. Ray 
florets neuter or female, sterile; limb yellow. Disc corolla 
5-lobed. Ray cypselas (sterile) flat, with 2 lateral and 1 
adaxial resin canal; pappus of c. 4 free or basally connate 



139 

scales. Disc cypselas 5-angled with 5 thin ribs covered with 
myxogenic cells; pappus absent. 

Distribution. SW Europe in Spain and Portugal. - Mono- 
typic. 

Lepidophorum, formerly a member of Anthemis because of 
its paleate receptacle, is difficult to place. Several authors 
have suggested a relationship to Leucanthemum mainly 
because of the habitual (foliage) similarities, though Lepi- 
dophorum does not have the specialized fruits of the Leucan- 
themum group of genera. 

Hading (1960) investigated the embryology of many 
Anthemis species, including A. repanda {= Lepidophorum 
repandum). Lepidophorum has the common {Polygonum) 
type of monosporic embryo sacs, while Anthemis proper has 
tetrasporic embryo sacs. Harling also described the cypselas 
of Lepidophorum in detail and discussed the relationships of 
the genus. He concluded that Lepidophorum should be 
recognized as a separate genus, not related to Anthemis but 
possibly distantly so to Coleostephus, i.e. the Leucanthemum 
group of genera. Lepidophorum is here provisionally placed 
as a basal member of subtribe Leucantheminae, where the 
Leucanthemum group is a subclade (Fig. 10, LeS). 

64. NIPPON ANTHEMUM Kitam. in Acta phytotax. 
geobot. Kyoto 29: 169 (1978). Type species: A^. 
nipponicum (Franchet ex Maxim.) Kitam. 

A shrub. Leaves alternate, obovate, apically serrate leaves 
rather densely set at the ends of the branches. Capitula rather 
large, solitary, pedunculate, radiate. Involucral bracts wide, 
many-veined. Receptacle convex, epaleate. Ray florets 
female, fertile; Umb white, many-veined. Disc corolla 
5-lobed. Cypselas oblong, 8-10-ribbed, rather thin-walled, 
without myxogenic cells. Pappus a corona of small scales. 

Distribution. Japan. - Monotypic. 

The monotypic Nipponanthemum is based on a handsome 
species formerly known as Chrysanthemum nipponicum 
Franchet ex Maxim. The position of Nipponanthemum is very 
unclear. It is provisionally placed as an aberrant member of 
subtribe Leucantheminae. It may on the other hand be more 
closely related to the relatively plesiomorphic members of 
Artemisiinae, i. e. Dendranthema or certain species in that 
genus. When he described the genus Kitamura considered it 
related to Argyranthemum but we find no support for his 
conclusion. 

65. LEUCANTHEMELLA Tzvelev in Komarov, Fl. 
URSS 26: 137 (1961). Type species: L. serotina (L.) 
Tzvelev. 

Perennial herbs. Leaves alternate, entire or serrate. Capitula 
solitary or laxly corymbose, pedunculate, radiate. Receptacle 
convex, epaleate. Ray florets female, sterile; hmb white or 
reddish, many-veined. Disc corolla 5-lobed, with a distinct 
enervate limb with sessile glands. Cypselas distinctly 7-12- 
ribbed, without myxogenic cells, with an apical rim but 
pappus absent. 

Distribution. E. Europe (L. serotina) and Far East, China 
in Manchuria, Korea and Japan (L. linearis). - 2 spp. 

This genus is difficult to place (see Dienst, 1983). Tzvelev 



140 



K. BREMER AND C. J. HUMPHRIES 



also noted its isolated position. In habit it is similar to other 
members of subtribe Leucantheminae, where it is provision- 
ally placed. It may also be related to Dendranthema or some 
species of that genus. Leucanthemella has distinctly many- 
ribbed, non-myxogenic fruits. This may be related to habitat; 
the two species grow in marshy places. 

L. linearis (Matsum.) Tzvelev 
L. serotina (L.) Tzvelev 



66. NIVELLEA Wilcox, Bremer & Humphries, gen. 
nov. Type species: N. nivellei (Braun-Blanquet & 
Maire) Wilcox, Bremer & Humphries. 

Herba annua. Folia lobata sublacerata. Capitula solitaria 
pedunculata radiata. Receptaculum epaleaceum. Flosculi 
radiati limbo albo. Flosculi disci tubo basaliter incrassato. 
Cypselae oblongae, 5-8-costata, sine canalibus secretoriis, 
sine celluHs mucilaginis. Pappus nuUus. 

An annual herb. Leaves lobed and rather lacerate. Capitula 
solitary, pedunculate, radiate. Receptacle flat to convex, 
epaleate. Ray florets female, fertile; limb white. Disc corolla 
5-lobed; tube basally swollen in fruit. Cypselas oblong, 
5-8-ribbed, without resin canals and myxogenic cells. Pappus 
absent. 

Distribution. N. Africa in Morocco. - Monotypic. 

The position of Nivellea is difficult to assess. It is superficially 
similar to some members of the Leucanthemum group, e. g. 
Leucoglossum, but does not have the specialized cypselas of 
that group. It is here provisionally placed as a basal member 
of subtribe Leucantheminae. 

N. nivellei (Braun-Blanquet & Maire) Wilcox, Bremer & 
Humphries, comb. nov. Basionym: Chrysanthemum nivellei 
Braun-Blanquet & Maire in Bull. Soc. Hist. nat. Afr. N. 13: 
187 (1922). 



67. PHALACROCARPUM (DC.) Willk. in Bot. Ztg 
22: 252 (1864). Type species: P. oppositifolium 
(Brot.) Willk. 

Creeping, suffruticose perennials. Leaves opposite, sheath- 
ing, serrate to pinnatifid. Capitula solitary, pedunculate, 
radiate. Involucral bracts with dark brown margins. Recep- 
tacle convex, epaleate. Ray florets female, fertile; limb white 
or purplish. Central disc florets male (style-branches fused) 
or neuter. Disc corolla 5-lobed. Cypselas 7-9-ribbed; testa 
epidermis thick-walled and dark reddish, of elongate-sinuate 
cells. Pappus absent. 

Distribution. SW Europe in Spain and Portugal. - 2 spp. 

Phalacrocarpum with opposite leaves is in habit and leaf 
shape similar to other members of Leucantheminae. The 
subtribal position of this genus must nevertheless be consid- 
ered provisional. 

P. hoffmannseggii (Samp.) Lainz 

P. oppositifolium (Brot.) Willk. (P. anomalum Cout.) 



68. LEUCANTHEMOPSIS (Giroux) Heyw. in An. 
Inst. bot. A. J. Cavanilles 32: 181 (1975). Type 
species: L. alpina (L.) Heyw. 

Creeping or caespitose suffruticose perennials. Leaves alter- 
nate, serrate to pinnatifid, generally pectinate and spathulate 
in outline. Capitula sohtary, rather long-pedunculate, radi- 
ate. Involucral bracts sometimes with dark brown margins. 
Receptacle convex, epaleate. Ray florets female, fertile; limb 
white to pinkish or yellow. Disc corolla 5-lobed; tube basally 
somewhat swollen in fruit. Cypselas 3-10- ribbed, with myxo- 
genic cells especially on the ribs, often with large 2-celled 
glands. Pappus a scarious flimsy corona. Flavonol 
5-glycosides present. 

Distribution. S. Europe mainly in Spain but extending to 
SW Russia (L. alpina), one species in N. Africa in Morocco 
(L. longipectinata). - 9 spp. 

Leucanthemopsis was originally a subsection of Tanacetum 
(sect. Pyrethrum), Tanacetum then taken in a very wide 
sense. Heywood (1954, 1976) noted its intermediate position 
between Tanacetum and Leucanthemum. Obviously, the Leu- 
canthemopsis species could not be accommodated in any of 
these two large genera, and the subsection was elevated to 
the rank of genus. New taxa have recently been described by 
Marchi (1980). 

In floral morphology and foliage Leucanthemopsis agrees 
with Hymenostemma, a possible sister group. Species of 
Leucanthemopsis are creeping or caespitose suffruticose 
perennials, whereas Hymenostemma is an annual herb. It 
seems to be the only reliable difference. On the other hand 
Hymenostemma groups with the specialized Prolongoa, both 
being annuals. The three genera clearly form a monophyletic 
group, but their interrelationships are uncertain. It is also 
possible that Leucanthemopsis is paraphyletic with Hymenos- 
temma and Prolongoa excluded. The matter requires further 
study. 

L. alpina (L.) Heyw. 

L. flaveola (Hoffsgg & Link) Heyw. 

L. longipectinata (Font Quer) Heyw. 

L. minima (Villars) Marchi 

L. pallida (Miller) Heyw. 

L. pectinata (L.) Lopez Gonzalez & Jarvis (L. radicans 

(Cav.) Heyw.) 
L. pulverulenta (Lagasca) Heyw. 
L. tatrae (Vierh.) Holub 
L. tomentosa (Lois.) Marchi 

69. HYMENOSTEMMA (Kunze) Willk. in Bot. Ztg 
22: 253 (1864). Type species: H. pseudanthemis 
(Kunze) Willk. - Prolongoa sect. Hymenostemma 
Kunze, pro parte. 

An annual herb. Leaves alternate, pinnatif id-pectinate, 
spathulate in outline. Capitula solitary, rather long- 
pedunculate, radiate. Receptacle convex to conical, epaleate. 
Ray florets female, sterile; limb white. Disc corolla 5-lobed; 
tube basally swollen in fruit. Cypselas 5-6-ribbed, with myxo- 
genic cells along the ribs. Pappus a scarious, flimsy corona. 

Distribution. SW Europe in Spain and N. Africa in 
Morocco. - Monotypic. 

Hymenostemma is very similar in habit to Prolongoa, though 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



141 



the latter is distinguished by a number of floral autapomor- 
phies. Hymenostemma differs from Leucanthemopsis by its 
annual habit, a feature shared by Prolongoa. The sister group 
relationships of these three genera are somewhat uncertain 
and it is even possible that Hymenostemma and Prolongoa, 
together or separately, have their sister groups within Leu- 
canthemopsis. 

70. PROLONGOA Boiss., Voyage hot. Espagne 2: 320 
(1840). Type species: P. hispanica Lopez Gonzalez 
& Jarvis. 

An annual herb. Leaves alternate, pinnatif id-pectinate, gen- 
erally spathulate in outline. Capitula soUtary, rather long- 
pedunculate, radiate. Receptacle convex, epaleate. Ray 
florets neuter; limb yellow; pappus a large flimsy corona. 
Disc corolla 5-lobed. Style-branches long-penicillate. Cypse- 
las with 1 adaxial and 2 lateral rather thick ribs and 2 abaxial 
ribs, with myxogenic cells along the ribs; cypsela wall with 
rod-shaped crystals in small packets; pappus absent. 

Distribution. SW Europe in Spain. - Monotypic. 

Prolongoa has been proposed and now accepted for conser- 
vation (Bremer et al., 1987). The species has been known 
erroneously as Prolongoa pectinata, a name that cannot be 
used since the basionym is a species of Leucanthemopsis. 
Prolongoa is related to Hymenostemma and Leucanthemop- 
sis. The matter is further discussed under these genera. 
Prolongoa has a number of autapomorphies, e. g. a reduced 
disc floret pappus and more strongly developed cypsela ribs. 

71. LEUCANTHEMUM Miller, Card. Diet. abr. 4th 
ed. (1754). Type species: L. vulgare Lam. 

Perennial herbs with red-tipped roots. Leaves alternate, 
entire, serrate, or pinnatifid. Capitula solitary or laxly corym- 
bose, pedunculate, radiate or discoid. Receptacle convex or 
sometimes conical, epaleate. Ray florets female, fertile; limb 
white, pink, or rarely yellow. Disc corolla 5-lobed; tube 
basally swollen and spongy in fruit, especially abaxially. 
Cypselas c. 10-ribbed with vallecular lacunae and vallecular 
secretory canals as well as vascular strands between the ribs, 
with myxogenic cells along the ribs. Pappus a corona or an 
adaxial auricle, sometimes absent. Flavonol 5-glycosides 
present. 

Distribution. Throughout Europe but mainly C. and S. 
parts, one species (L. discoideum) also in N. Africa in 
Morocco, Algeria, and Tunisia, some species (e.g. L. vul- 
gare) widespread as weeds. - 33 spp. 

With the removal of L. paludosum {=Leucoglossum palu- 
dosum) and L. arundanum {=Rhodanthemum arundanum) 
as well as the North African Leucanthemum species to 
Leucoglossum and Rhodanthemum, Leucanthemum becomes 
morphologically homogeneous and defined by its anthocya- 
nin red root tips. The latter are present in all herbaceous 
perennials (Favarger, 1966) which now constitute the genus 
Leucanthemum s. s. The genus is a polyploid complex (Vil- 
lard, 1970). 

L. adustum (Koch) Gremli 
L. aligulatum Vogt 
L. atratum (Jacq.) DC. 
L. burnatii Briq. & Cav. 



L. catalaunicum Vogt 

L. chloroticum A. Kerner & Murb. 

L. corsicum (Less.) DC. 

L. crassifolium (Lange) Willk. in Willk. & Lange 

L. cuneifolium Le Grand ex Coste 

L. delarbrei Timb.-Lagr. 

L. discoideum (All.) Coste (L. fontanesii Boiss. & Reuter, 

Chrysanthemum fontanesii (Boiss. & Reuter) Quezel & 

Santa) 
L. favargeri Vogt 
L. gaudinii Dalla Torre 
L. gracilicaule (Duf.) Alavi & Heyw. 
L. graminifolium (L.) Lam. 
L. heterophyllum (Willd.) DC. 
L. ircutianum DC. 
L. laciniatum Huter, Porta & Rigo 
L. lacustre (Brot.) Samp. 
L. leucolepis (Briq. & Cav.) Horvatic 
L. maestracense Vogt & Hellwig 
L. maximum (Ram.) DC. 
L. meridionale Le Grand 
L. merinoi Vogt & Castroviejo 
L. monspeliense (L.) Coste 
L. montserratianum Vogt 
L. pallens (Gay in Perreymond) DC. 
L. praecox (Horvatic) Horvatic 
L. pluriflorum Paul. 
L. subglaucum De Laramb. 
L. sylvaticum (Brot.) Nyman 
L. vulgare Lam. 
L. waldensteinii (Schultz-Bip.) Pouzar 

72. RHODANTHEMUM Wilcox, Bremer & 
Humphries, comb, et stat. nov. Type species: R. 
arundanum (Giroux) Wilcox, Bremer & Humphries 
{Leucanthemum sect. Rhodanthemum Vogt, 
Leucanthemum subg. Chrysanthemopsis Maire, 
nom. nud.). 

Plantae perennes stolonibus saepe tegetes formantes. Folia 
alterna aggregata rosulata trifida vel ternato-pinnata ut vide- 
tur longi-petiolata. Capitula solitaria longepedunculata 
radiata. Bracteae involucri margine atro-fuscae. Receptacu- 
lum convexum epaleaceum. Flosculi radii feminei, vel fertiles 
vel steriles, limbo albido vel roseo vel rubro vel cremeo- 
aurantici multi-venoso. Flosculi disci corolla quinquiloba, 
lutea vel rubra tubo basi inflato et spongioso in fructus 
maturitate. Cypselae 5-12-costae valleculis lacunis et canali- 
culis secretoriis cum fills vascularibus et cellulis myxogenis 
intercostalibus instructis; costae protusae angustae et aliquan- 
tum aliformes. Pappus e corona scariosa vel auricula adaxiali 
basi coroniformi sistens. 

Stoloniferous and often mat-forming perennials. Leaves 
alternate, closely set, becoming rosulate, trifid or ternate- 
pinnafifid and seemingly long-petiolate. Capitula solitary, 
with long peduncles, radiate. Involucral bracts with dark 
brown margins. Receptacle convex, epaleate. Ray florets 
female, fertile or sterile; limb white, pink, reddish, or creamy 
orange, many-veined. Disc corolla 5-lobed, yellow or red; 
tube basally swollen and spongy at maturity of the fruit. 
Cypselas 5-12-ribbed with vallecular lacunae and vallecular 
secretory canals as well as vascular strands between the ribs, 
and with myxogenic cells along the ribs; ribs protruding. 



142 



K. BREMER AND C. J. HUMPHRIES 



narrow and somewhat wing-shaped. Pappus with a scarious 
corona or an adaxial basally coroniform auricle. 

Distribution. N. Africa in Morocco and Algeria, one spe- 
cies (R . arundanum) also in SW Europe in Spain. - 12 spp. 

Rhodanthemum (see Vogt, 1991) is a well characterized 
group of North African, Atlas montane perennials which 
form mats at relatively high altitudes (700-1200 m). They 
were formerly classified in Leucanthemum or Chrysanthe- 
mum but are distinguished by a number of synapomorphies. 

R. arundanum (Boiss.) Wilcox, Bremer &. Humphries, comb, 
nov. Basionym: Pyrethrum arundanum Boiss., Voy. hot. 
Espagne 2: 317 (1840) {Leucanthemum arundanum (Boiss.) 
Cuatrec, Leucanthemum mairei Humbert). 

R. atlanticum (Ball) Wilcox, Bremer & Humphries, comb, 
nov. Basionym: Chrysanthemum atlanticum Ball in/. Bot., 
Land. 11: 366 (1873) {Leucanthemum atlanticum (Ball) 
Maire). 

R. briquetii (Maire) Wilcox, Bremer & Humphries, comb, 
nov. Basionym: Leucanthemum briquetii Maire in Bull. 
Soc. Hist. nat. Afr. N. 15: 88 (1924). 

R. catananche (Ball) Wilcox, Bremer & Humphries, comb, 
nov. Basionym: Chrysanthemum catananche Ball in J. Bot. 
Lond. 11: 366 (1873) {Leucanthemum catananche (Ball) 
Maire). 

R. depressum (Ball) Wilcox, Bremer & Humphries, comb, 
nov. Basionym: Chrysanthemum gayanum var. depressum 
Ball in J. Linn. Soc. 16: 509 (1878) {Leucanthemum 
depressum (Ball) Maire). 

R. gayanum (Cosson & Durieu) Wilcox, Bremer & 
Humphries, comb. nov. Basionym: Pyrethrum gayanum 
Cosson & Durieu in Bull. Soc. bot. Fr. 4: 15 (1857) 
{Chrysanthemum gayanum Ball, Leucanthemum gayanum 
(Cosson & Durieu) Maire). 

R. hosmariense (Ball) Wilcox, Bremer & Humphries, comb, 
nov. Basionym: Chrysanthemum maresii var. hosmariense 
Ball in J. Bot. Lond. 11: 366 (1873) {Leucanthemum 
hosmariense (Ball) Font Quer). 

R. maresii (Cosson) Wilcox, Bremer & Humphries, comb, 
nov. Basionym: Pyrethrum maresii Cosson in Bull. Soc. 
bot. Fr. 4: 16 (1857) {Leucanthemum maresii (Cosson) 
Maire). 

R. maroccanum (Battand.) Wilcox, Bremer & Humphries, 
comb. nov. Basionym: {Chrysanthemum maroccanum 
Battand. in Bull. Soc. Hist. nat. Afr. N. 12: 189 (1921) 
{Leucanthemum maroccanum (Battand.) Maire). 

R. mesatlanticum (Emb. & Maire) Wilcox, Bremer & 
Humphries, comb. nov. Basionym: Leucanthemum mesat- 
lanticum Emb. & Maire, PI. Maroc Nov. (Arch. Sci. 
Maroc.) Fasc. II: 5 (1929). 

R. pseudo-catananche (Maire) Wilcox, Bremer & 
Humphries, comb. nov. Basionym: Leucanthemum pseudo- 
catananche Maire in Mem. Soc. Sci. nat. Phys. Maroc 15:37 
(1926). 

R. redieri (Maire) Wilcox, Bremer & Humphries, comb. nov. 
Basionym: Leucanthemum redieri Maire in Mem. Soc. Sci. 
nat. Phys. Maroc 15:38 (1926). 



73. LEUCOGLOSSUM Wilcox, Bremer & 
Humphries, gen. nov. Type species: L. paludosum 
(Poiret) Wilcox, Bremer & Humphries - Prolongoa 
sect. Hymenostemma Kunze pro parte. 

Herbae annuae. Folia dentata-serrata vel pinnatifida. 
Capitula solitaria radiata. Receptaculum epaleaceum. Flos- 
culi radiati limbo albo vel flavido, basaliter luteo; pappus 
coroniformis scariosus vel adaxialiter auriculiformis. Flosculi 
disci corolla parum zygomorpha tubo incrassato; pappus 
plerumque nullus. Cypselae ellipsoideae parvae 7-10- 
costatae, inter costas lacunis vallecularibus canalibus secre- 
toriis et fasciculis vascularibus, in costis cellulis mucilaginis 
instructae. 

Annual herbs. Leaves alternate, dentate-serrate to pinnatifid. 
Capitula solitary, pedunculate, radiate. Receptacle convex or 
conical, epaleate. Ray florets female, fertile; limb white or 
pale yellow with a yellowish base; pappus a scarious corona 
or an adaxial auricle. Disc corolla 5-lobed, slightly zygomor- 
phic; tube basally swollen in fruit. Cypselas ellipsoid, com- 
paratively small, c. 1 mm long, 7-10-ribbed with vallecular 
lacunae and vallecular secretory canals as well as vascular 
strands between the ribs, with myxogenic cells along the ribs; 
generally the pappus is absent. 

Distribution. SW Europe (L. paludosum) in Spain and N. 
Africa (all species) in Morocco, Algeria, Tunisia, and Libya. 
- 3 spp. 

The type species of this new genus has been recognized as 
anomalous within Leucanthemum e. g. in Flora europaea 
(Tutin et al., 1976). It differs by its annual habit, the small 
cypselas and the pappus, present in ray florets but absent in 
disc florets. Together with two North African species it is 
more closely related to Chlamydophora, another annual with 
small cypselas. 

Leucoglossum paludosum is often cultivated as an orna- 
mental. 

L. decipiens (Pomel) Wilcox, Bremer & Humphries, comb. 

nov. Basionym: Leucanthemum decipiens Pomel, Nouv. 

mat.fl. ad.: 59 {I860). 
L. paludosum (Poiret) Wilcox, Bremer & Humphries, comb. 

nov. Basionym: Chrysanthemum paludosum Poiret, Voy. 

Barbaric 2: 241 (1789). {Leucanthemum paludosum 

(Poiret) Bonnet & Barratte, Leucanthemum setabense 

DC, Hymenostemma paludosum (Poiret) Pomel). 
L. reboudianum (Pomel) Wilcox, Bremer & Humphries, 

comb. nov. Basionym: Leucanthemum reboudianum 

Pomel, Nouv. mat. fl. ad.: 291 (1860). 

74. CHLAMYDOPHORA Ehrenb. ex Less., Syn. gen. 
Compos.: 265 (1832). Type species: C. tridentata 
(Dei.) Ehrenb. ex Less. 

An annual herb. Leaves alternate or basally opposite, entire 
or tridentate, somewhat fleshy. Capitula solitary, peduncu- 
late, discoid. Receptacle convex, epaleate. Corolla 4- or 
5-lobed, yellow or sometimes reddish; tube basally swollen 
and spongy in fruit. Cypselas ellipsoid, comparatively small, 
c. 1 mm long, 6-10-ribbed with vallecular lacunae and 
vallecular secretory canals as well as vascular strands between 
the ribs, with myxogenic cells on the ribs. Pappus a large, 
scarious, adaxial but basally coroniform auricle, as long as the 
corolla or longer. 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 

Distribution. N. Africa in Tunisia, Libya, and Egypt, SE 
Europe in Greece and in Cyprus. - Monotypic. 

Chlamydophora is the monotypic and discoid sister group of 
Leucoglossum. The former C. pubescens is transferred to 
Aaronsohnia. C. tridentata is known also as Tripleurosper- 
mum tridentatum Hoffm. 

75. CHRYSANTHOGLOSSUM Wilcox, Bremer & 
Humphries, gen. nov. Type species: C. trifurcatum 
(Desf.) Wilcox, Bremer & Humphries. 

Herbae annuae vel raro biennes. Folia pinnatisecta vel saepe 
trifurcata. Capitula solitaria, pedunculata, radiata. Bracteae 
involucri latae, plurinerves, flabelliformes. Receptaculum 
epaleaceum. Flosculi radiati limbo aureo; pappus adaxialiter 
auriculiformis scariosus vel nullus. Flosculi disci tubo incras- 
sato spongioso in fructu; pappus coroniformis rigidus extus 
cellulis mucilaginis instructus. Cypselae c. 10-costata, inter 
costis lacunis vallecularibus canalibus secretoriis et fasciculis 
vascularibus, in costis et corona cellulis mucilaginis instruc- 
tae. 

Annual or rarely biennial herbs. Leaves alternate, pinnati- 
sect, often trifurcate. Capitula solitary, pedunculate, radiate. 
Involucral bracts wide, many-veined, flabelliform. Recep- 
tacle convex, epaleate. Ray florets female, sterile or some- 
times fertile; limb golden yellow, many-veined; pappus a 
scarious adaxial auricle or absent. Disc corolla 5-lobed; tube 
basally swollen and spongy in fruit; lobes with appendages; 
pappus a rather stiff corona with myxogenic cells on the 
outside. Cypselas dorsiventrally somewhat compressed, c. 
10-ribbed with vallecular lacunae and vallecular secretory 
canals as well as vascular strands between the ribs, with 
myxogenic cells along the ribs and also on the pappus. 

Distribution. N. Africa in Morocco, Algeria, Tunisia and 
Libya. - 2 spp. 

Both species are readily recognizable as sister species by the 
distinctive leaves and cypselas. Chrysanthoglossum is the 
sister group of the three genera Glossopappus, Coleostephus, 
and Plagius. 

C. deserticola (Murb.) Wilcox, Bremer & Humphries, comb. 

nov. Basionym: Pyrethrum deserticola Murb., mActa Univ. 

lund. 33(12): 98 (1897) {Chrysanthemum deserticola 

(Murb.) F. Buxbaum). 
C. trifurcatum (Desf.) Wilcox, Bremer & Humphries, comb. 

nov. Basionym: Chrysanthemum trifurcatum Desf., Fl. 

atlant. 2: 281 (1799) {Leucanthemopsis trifurcata (Desf.) 

Alavi). 



143 

bulbous callus; pappus a large, scarious, adaxial, flabelliform 
auricle, as long as the corolla or longer. 

Distribution. SW Europe in Spain and Portugal, N. Africa 
in Morocco, Algeria and Tunisia. - Monotypic. 

Glossopappus macrotus is related to Coleostephus and 
Plagius. Possibly Glossopappus has its sister group within a 
paraphyletic Coleostephus. The three genera form a mono- 
phyletic group. They are further discussed under Coleoste- 
phus. 

11. COLEOSTEPHUS Cass, in Diet. Sci. Nat. 41: 43 
(1826). Type species: C. myconis (L.) Reichenb. f. 

Annual herbs. Leaves alternate, serrate-dentate, spathulate. 
Capitula solitary or laxly corymbose, pedunculate, radiate. 
Involucral bracts wide, more or less flabelliform. Receptacle 
convex to conical, epaleate. Ray florets female, fertile; limb 
golden yellow, many-veined. Disc corolla 5-lobed; tube 
basally much swollen and spongy in fruit, especially abaxially; 
lobes with more or less developed appendages. Cypselas 
arcuate, 8-10-ribbed with vallecular lacunae and vallecular 
secretory canals as well as vascular strands between the ribs, 
with myxogenic cells on the ribs; ribs basally and adaxially 
fused into a bulbous callus; pappus an oblique, adaxially 
more or less strongly developed scarious corona. Flavonol 
5-glycosides present. 

Distribution. S. Europe and N. Africa from Morocco to 
Libya. - 3 spp. 

Coleostephus is related to Plagius and Glossopappus (Alavi, 
1968, 1988; Alavi & Heywood, 1976). The six species of these 
three genera form a monophyletic group but their interrela- 
tionships are unclear. Plagius with two species is diagnosed 
by being discoid and perennial (probably secondarily) but 
these characters are homoplasious and not supported by 
stronger synapomorphies. The monotypic Glossopappus has 
a specialized large pappus auricle. There is no synapomorphy 
to diagnose Coleostephus from the other two genera and it is 
possibly paraphyletic. Coleostephus and Plagius appear to be 
related by their more arcuate cypselas but the character is 
variable within Coleostephus. 

C. multicaulis (Desf.) Durieu {Chrysanthemum multicaule 
Desf.) 

C. myconis (L.) Reichenb. f. {Chrysanthemum myconis L., 
Kremeria myconis (L.) Maire) 

C. paludosus (Durieu) Alavi (C. clausonis Pomel, Chrysan- 
themum clausonis (Pomel) Battand., Kremeria paludosa 
Durieu) 



76. GLOSSOPAPPUS Kunze in Flora, Jena 29: 748 
(1846). Type species: G. chrysanthemoides Kunze 
(G. macrotus (Durieu) Briq. in Burnat). 

An annual herb. Leaves alternate, serrate-dentate, obovate- 
spathulate. Capitula solitary, pedunculate, radiate. Involu- 
cral bracts wide, flabelliform. Receptacle conical, epaleate. 
Ray florets female, fertile; limb golden yellow, many-veined. 
Disc corolla 5-lobed; tube basally much swollen and spongy 
in fruit, especially abaxially; lobes with appendages. Cypselas 
somewhat arcuate, 8-10-ribbed with vallecular lacunae and 
vallecular secretory canals as well as vascular strands between 
the ribs, with myxogenic cells along the ribs; basally with a 



78. PLAGIUS L'Herit. ex DC, Prodr. 6: 135 (1838). 
Type species: P. ageratifolius (Desf.) L'Herit. ex 
DC. {P. flosculosus (L.) Alavi & Heyw.). 

Herbaceous or suffruticose perennials. Leaves alternate, 
serrate-dentate, spathulate. Capitula solitary or laxly corym- 
bose, pedunculate, discoid. Receptacle flat or convex, epale- 
ate. Disc corolla 5-lobed; tube basally swollen and spongy in 
fruit; lobes with appendages. Cypselas arcuate, c. 10-ribbed, 
with vallecular lacunae and vallecular secretory canals as well 
as vascular strands between the ribs; basally with a bulbous 
callus, with myxogenic cells on the ribs. Pappus an adaxial 
auricle. Flavonol 5-glycosides present. 



144 



K. BREMER AND C. J. HUMPHRIES 



Distribution. S. Europe in Corsica and Sardinia (P. floscu- 
losus) and N. Africa in Algeria and Tunisia {P. grandis). - 2 
spp. 

The two species are very different from each other. Alavi & 
Heywood (1976) united them into one genus because both 
are discoid. P. grandis is instantly recognizable by the larger 
solitary capitula on unbranched, basally hairy stems. P. 
flosculosus, by contrast, has smaller capitula on branched 
stems and it is similar to species of the related genera 
Coleostephus and Glossopappus, except for the discoid 
capitula. Plagius, Coleostephus, and Glossopappus form a 
monophyletic group but their interrelationships deserve fur- 
ther study (see under Coleostephus). The species known as P. 
virgatus (Desf.) DC. is synonymous with Leucanthemum 
discoideum, following Alavi & Heywood (1976). 

P. flosculosus (L.) Alavi & Heyw. 

P. grandis (L.) Alavi & Heyw. {Chrysanthemum grandiflo- 
rum (Desf.) Battand.) 

11. THAMINOPHYLLINAE Bremer & 
Humphries, subtrib. nov. 

Type species: Thaminophyllum multiflorum Harvey 

Herbae perennes vel suffrutices vel frutices. Folia serrata vel 
dentata vel Integra vel solum pauciloba. Capitula solitaria, 
radiata. Cypselae oblongae vel ellipsoideae. Pappus coroni- 
formis vel nullus. 

Perennial herbs, half-shrubs or shrubs. Leaves serrate, den- 
tate, entire, or few-lobed only. Capitula solitary, radiate. 
Receptacle flat to conical, paleate or epaleate, sometimes 
pilose. Ray floret limb white or yellow. Disc corolla 4- or 
5-lobed. Cypselas oblong to ellipsoid, 3-8-ribbed (or angled) 
or rarely 10-ribbed with costal veins and resin canals {Ade- 
nanthellum), without or rarely with large myxogenic cells in 
scattered groups {Thaminophyllum). Pappus a corona or 
absent. 

Distribution (Table 22). S. Africa. - 5 genera, 17 spp. 

Table 22 General distribution of Thaminophyllinae and genera. 
x=indigenous, o=introduced. 

S. Afr. 



Thaminophyllinae 


X 


Osmitopsis 


X 


Adenanthellum 


X 


Inezia 


X 


Lidbeckia 


X 


Thaminophyllum 


X 



This subtribe comprises some small South African genera, 
which have received little attention in general treatments of 
the Anthemideae. Thaminophyllum, Lidbeckia, Inezia, and 
Adenanthellum are related as noted by earlier authors. Bond 
(1980) and Goldblatt (1980) stated that Thaminophyllum is 
related to Lidbeckia and Inezia. The unusual chromosome 
number x=10, known in Thaminophyllum and Inezia, is a 
feature mentioned by Bond and Goldblatt. Inezia was origi- 
nally described as a Lidbeckia. Adenanthellum was assumed 
to be most closely related to Inezia by Nordenstam (1976) 
when he originally described the genus. 



Osmitopsis was earlier classified in Inuleae because of the 
tailed anthers and its proper relationship within Anthemideae 
has hitherto not been considered. A preliminary chromosome 
count of 2n=10 as well as remarkable similarities between 
some species of Osmitopsis and the other genera indicate that 
these genera are related, and it is probable that together they 
form a monophyletic group. The cladogram is the single most 
parsimonious one derivable from the data matrix. 

Clades and characters - Fig. 11, Tables 2, 23. 



|=Thl: 



7= 79 Osmitopsis 



Th3 



Jc 



it=Th2 



f r- 

iTh4JC 



80 Adenanthellum 

81 Inezia 

82 Lidbeckia 

83 Thaminophyllum 



Fig. 11 Cladogram of the Thaminophyllinae produced by the ie 
option in Hennig86. Cladogram length = 25, consistency index = 
92, retention index = 80. 

Table 23 Data matrix for the Thaminophyllinae. 1 = presence, = 
absence, ? = missing data or not applicable, p = polymorphic but 
scored as the plesiomorphic condition, a = polymorphic but scored 
as the apomorphic condition. 

nil 1 1111 11 1 nil 

145057652 745966 5892337574081 24 177675 
532718111 75620157784132928538262 365600 

79. Osmitopsis allll?10a ini000000000p000p?0p00 ???00p 

80. Adenanthellum allll?lll ?000111in 1111000070000 ?????? 

81. Inezia alOn?101 100011 11000000110070000 77700? 

82.Lidbeckia 111117100 70001100000001011111000 ?????7 

8?>.ThaminophyllumOnni\m 10001100000001011110111 777777 



Clade Thl - subtribe Thaminophyllinae 

51 reversed. Floral parts without resin canals. Floral resin 
canals are present in a number of subtribes and also in 
Thaminophyllinae, although only in Adenanthellum. This is 
most parsimoniously explained as a reversal for the subtribe 
with a reappearance in Adenanthellum. Clearly, the character 
is difficult to interpret. 

177 Chromosome number x=IO. The chromosome number is 
unknown in Adenanthellum and Lidbeckia, but is neverthe- 
less best interpreted as a synapomorphy for the whole sub- 
tribe. 

Osmitopsis 

45 Receptacle paleate. 

56 Ray floret limb epidermis cells tabular (senecioid or muti- 
sioid type). This ligule epidermis type is uncommon in 
Anthemideae and reported for example in Osmitopsis 
(Baag0e, 1977). 

92 Anthers caudate. 

Clade Th2 

60 Ray floret tube confluent with the cypsela. 

61 Ray floret tube persistent on the cypsela. In Adenanthellum 
and Inezia the tube sinus extends to the base and the tube is 
then virtually absent. Nevertheless the ray corolla, in these 
cases the limb, is both confluent with and persistent on the 
cypsela. 

Clade Th3 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



145 



5 Plants with one or few sparsely branched stems arising from 
a woody villous caudex. Some species of Osmitopsis are 
similar to Adenanthellum and Inezia in this aspect, as men- 
tioned by Nordenstam (1976) when grouping the latter two 
genera together. 

57 Ray floret tube sinus extending to the base. 

Adenanthellum 

51 Floral parts with resin canals. See note under clade Thl. 
87 Disc corolla tube confluent with the cypsela. 

98 Pollen grains hexa-panto-colporate. 

124 Cypselas with 10 (8-12) multicellular epicarpic ribs. 

.131 Cypselas 10-ribbed with costal veins and resin canals. 

133 Cypselas with costal resin canals or sacs. This is character- 
istic, for example, of Lepidophorum in Leucantheminae and 
large part of Matricariinae but in Adenanthellum the cypselas 
are also 10-ribbed. Whether the resin canals are homologous 
in the two groups or not is difficult to assess. It is interpreted 
here as a parallelism. 

172 Pappus absent in ray and disc cypselas. 

Inezia 

52 reversed. Ray floret limb yellow not white. 

59 Ray floret tube and cypsela pilose laterally; ray floret limb 
pilose abaxially. This character also occurs in one of the two 
Lidbeckia species (L. pectinata). 

72 Disc corolla 4-lobed. 

Clade Th4 

21 reversed. Leaves not serrate-dentate. 

48 Receptacle pilose. 

72 Disc corolla 4-lobed. 

105 Stylopodium large and persistent in fruit. This character 
occurs also in some species of Osmitopsis. 

172 Pappus absent in ray and disc cypselas. 

183 Dehydrofalcarinone and dehydrofalcarinol present. The 
other genera of this subtribe are unknown chemically. 

Lidbeckia 

18 Leaves with few, oblong to rounded, apically mucronate 
lobes. 

Thaminophyllum 

2 Plants shrubby. 

15 reversed. Leaves entire, not variously deeply lobed or 
divided. 

26 Leaves closely set, lanceolate to linear. 

142 Cypselas with large myxogenic cells in rounded, scattered 
groups. 



79. OSMITOPSIS Cass, in Bull. Sci. Soc. philom. 
Paris 1817: 154 (1817). Type species: O. 
asteriscoides (P. Bergius) Less. 

Shrubs or half-shrubs. Leaves alternate, lobed, dentate, 
serrate, or entire, often glandular-punctate. Capitula solitary 
or laxly corymbose, radiate. Receptacle flat or convex to 
conical, paleate. Ray florets female, fertile or sterile, or 
neuter; tube occasionally pilose; limb white, many-veined, 
occasionally abaxially pilose; epidermis cells tabular. Disc 
corolla 5-lobed, glandular. Anthers caudate. Stylopodium 
sometimes large and persistent in fruit. Cypselas somewhat 
3-4-angled or -ribbed. Pappus a corona of subulate to trian- 
gular scales, or absent. 

Distribution. S. Africa in the Cape. - 9 spp. 

Until recently Osmitopsis has been placed in the Inuleae 
because of its tailed anthers. The traditional concept of the 
Anthemideae comprised genera without anther tails, a condi- 
tion which must be interpreted as a symplesiomorphy, how- 
ever. Tailed anthers have evolved independently within 
Anthemideae in the relatively unrelated genera Osmitopsis, 
Inulanthera (Gonosperminae), and Hippolytia (Tanacetii- 
nae). Stix (1960) showed that Osmitopsis has anthemoid 
pollen and the tribal position in the Anthemideae was con- 
firmed by Bremer (1972) and Reitbrecht (1974). 

The position of Osmitopsis within Anthemideae has hith- 
erto remained unclear (Heywood & Humphries, 1977). It is 
placed here in Thaminophyllinae with other small South 
African genera with similar habit, foliage, many-veined rays, 
and the less usual chromosome number of x=10. A prelimi- 
nary count of 2n=20 was made by Bremer (unpubl., on O. 
pinnatifida) but it needs confirmation. Various but not all 
species of Osmitopsis are similar to other genera of subtribe 
Thaminophyllinae in a number of characters, particularly in 
habit, pilose and sterile rays, the loss of pappus and a large 
stylopodium. The genus was revised by Bremer (1972, 1976). 

O. afra (L.) Bremer 

O. asteriscoides (P. Bergius) Less. 

O. dentata (Thunb.) Bremer 

O. glabra Bremer 

O. nana Schltr 

O. osmitoides (Less.) Bremer 

O. parvifolia (DC.) Hofmeyr 

O. pinnatifida (DC.) Bremer 

O. tenuis Bremer 

80. ADENANTHELLUM B. Nord. in Bot. Notiser 
132: 160 (1979). Type species: A. osmitoides 
(Harvey) B. Nord. - Adenanthemum B. Nord. 

A perennial herb with erect stems from a subterranean 
caudex. Leaves alternate, serrate, glandular-punctate. 
Capitula solitary, radiate. Receptacle convex, epaleate. Ray 
florets female, fertile; limb white, many-veined, confluent 
with the cypsela; tube absent (sinus extended to the base). 
Disc corolla 5-lobed, confluent with the cypsela. Pollen 
hexa-panto-colporate. Cypselas oblong, 10-ribbed, with 10 
veins and 10 resin canals. Pappus absent. 

Distribution. S. Africa in Natal and Transvaal and in 
Swaziland. - Monotypic. 

This recently described monotypic genus has a number of 



146 



K. BREMER AND C. J. HUMPHRIES 



autapomorphies, notably the unique pollen and the 10-ribbed 
cypselas with 10 resin canals. In habit and life-form, with a 
subterranean caudex generating herbaceous flowering stems, 
it is similar to Inezia, as indicated also by Nordenstam (1976) 
in his original description. 

A chromosome number of 2n=30 was reported by Goldb- 
latt (1980) who cited a preliminary, unpublished count by 
Nordenstam. From that one count (Nordenstam, pers. 
comm.) it is impossible to tell whether it represents an 
occasional triploid cell or a triploid specimen or taxon. 

The name Adenanthellum (Nordenstam, 1979) replaces 
Adenanthemum (Nordenstam, 1976), which turned out to be 
illegitimate. 

81. INEZIA E. Phillips in Bull. misc. Inf. R. bot. 
Gdns, Kew. 297 (1932). Type species: /. integrifolia 
(Klatt) E. Phillips. 

Perennial herbs with erect stems from a subterranean caudex. 
Leaves alternate, entire, glandular-punctate. Capitula soli- 
tary, radiate. Receptacle convex, epaleate. Ray florets 
female, fertile; limb yellow, many-veined, abaxially and 
basally laterally pilose, confluent with the laterally pilose 
cypsela; tube absent. Disc corolla 4-lobed, glandular. Cypse- 
las oblong, 4-angled. Pappus of minute scales. 

Distribution. S. Africa in Transvaal and in Swaziland. - 2 
spp. 

Inezia is based on the transfer of /. integrifolia from the 
related genus Lidbeckia. Brusse (1989fl) has shown that the 
two genera differ by several floral characters. Inezia has a 
stylopodium of thick-walled cells as compared with thin- 
walled cells in Lidbeckia and the nectaries are conspicuously 
larger in Lidbeckia. Also, there are 9-10 rows of cells in the 
filament collars of Lidbeckia as compared with 5-8 rows of 
cells in Inezia. Both genera have a 4-lobed corolla but 
nevertheless the sister group of Inezia is taken here to be 
Adenanthellum (with a 5-lobed corolla). These two genera 
have subterranean caudices from which herbaceous stems 
emerge (Nordenstam, 1976). They also have rays without a 
tube. It is more parsimonious to place Adenanthellum rather 
than Lidbeckia (as well as Thaminophyllum) as the sister 
group of this genus. 

/. integrifolia (Klatt) E. Phillips 
/. speciosa Brusse 

82. LIDBECKIA P. Bergius, Descr. pi. Cap.: 307 
(1767). Type species: L. pectinata P. Bergius 

Half-shrubs. Leaves alternate, glandular-punctate with few, 
oblong to rounded, apically mucronate lobes. Capitula soli- 
tary, rather long-pedunculate, radiate. Receptacle convex, 
epaleate, pilose. Ray florets female, sterile, or neuter; limb 
white, many- veined; sometimes dorsally pilose; tube some- 
times laterally pilose, confluent with the sometimes laterally 
pilose cypsela. Disc corolla 4-lobed, glandular, sometimes 
pilose; lobes sometimes with short acute appendages. Sty- 
lopodium large and persistent in fruit. Cypselas ellipsoid, 
3-8-ribbed. Pappus absent. Dehydrofalcarinone and dehy- 
drofalcarinol present. 

Distribution. S. Africa in the Cape. - 2 spp. 

The two species of Lidbeckia are related to Inezia and 



Thaminophyllum. They are held together as a pair by their 
similar foliage and habit. L. quinqueloba is more pubescent 
with pilose rays as in Inezia. On the other hand both species 
share a number of synapomorphies, for example the pilose 
receptacle, with Thaminophyllum. 

L. pectinata P. Bergius 

L. quinqueloba (L. f.) Cass. (L. lobata Thunb.) 

83. THAMINOPHYLLUM Harvey, Fl. cap. 3: 155 
(1865). Type species: T. multiflorum Harvey 

Shrublets. Leaves alternate, closely set, entire, lanceolate to 
linear, cricoid, glandular. Capitula solitary or laxly corym- 
bose, radiate. Receptacle convex or conical, epaleate, pilose. 
Ray florets female, sterile; limb white to somewhat purple- 
pink, many-veined; tube confluent with the cypsela. Disc 
corolla 4-lobed, glandular; lobes with acute or rounded 
appendages. Stylopodium large and persistent in fruit. Cypse- 
las ellipsoid, 3-4-angled, with large myxogenic cells in 
rounded, scattered groups. Pappus absent. Dehydrofalcar- 
inone and dehydrofalcarinol present. 

Distribution. S. Africa in the Cape. - 3 spp. 

Thaminophyllum is a small and distinct genus of cricoid 
South African shrublets. The sister group is Lidbeckia. The 
two genera differ much in foliage but have a similar floral 
morphology and they both have a pilose receptacle. The 
myxogenic cells on the cypselas are of a type different from 
the common Anthemideae pattern (of rows of large myxo- 
genic cells). In Thaminophyllum the myxogenic cells are 
arranged in rounded groups, scattered on and hardly elevated 
above the epicarp. Thaminophyllum was revised by Bond 
(1980). 

T. latifolium Bond 
T. multiflorum Harvey 
T. mundtii Harvey 

12. MATRICARIINAE Bremer & Humphries, 
subtrib. no v. 

Type species: Matricaria recutita L. 

Herbae annuae vel perennes vel suffrutices vel frutices. 
Capitula saepe solitaria vel raro corymbosa. Receptaculum 
epaleaceum vel raro paleaceum. Cypselae plerumque 5(4-6)- 
costatae costis varie dispositis vel interdum dorsiventraliter 
compressae et lateraliter alatae, plerumque abaxialiter et in 
costis sed non adaxialiter cellulis mucilaginis instructae. Pap- 
pus plerumque adaxiaHter longior, coroniformis vel auriculi- 
formis vel e squamis discretis compositus vel nullus. 

Annual or perennial herbs, shrublets or shrubs. Leaves often 
much dissected, sometimes few-lobed or entire or cricoid. 
Capitula solitary or rarely corymbose, pedunculate or rarely 
sessile, radiate, disciform or discoid. Receptacle variously 
shaped, epaleate or rarely paleate. Ray floret limb white or 
yellow. Outer female florets (in disciform capitula) in one or 
more rows, sometimes stalked. Disc or central florets 5- or 4-, 
rarely 3-lobed. Cypselas generally 5(4-6)-ribbed with differ- 
ent rib arrangements or sometimes dorsiventrally flattened 
and laterally winged, generally with myxogenic cells abaxially 
and on the ribs but not adaxially. Pappus generally adaxially 
long, a corona, an auricle, of separate scales, or absent. 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 

Distribution (Table 24). Worldwide, most genera in the 
Mediterranean region and in S. Africa, some Matricaria, 
Tripleurospermum, Cotula and Soliva species widespread as 
weeds. - 25 genera, 252 spp. 



147 

the absence of synapomorphies with genera of other subtribes 
they are provisionally retained in Matricariinae. The species 
of those two genera were formerly classified in Pentzia and 
Tripleurospermum. Rennera and Oncosiphon appear as sister 



Table 24 General distribution of Matricariinae and genera. x=indigenous, o=introduced. 



N.Am. 



Eur- 
Asia 



C.&E. 

Asia 



SW 
Asia 



S.Eur. N.Afr. S.Afr. 



Austr. 
N.Zeal. 



S.Am. 



Matricariinae 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Cymbopappus 














X 






Pentzia 












X 


X 






Marasmodes 














X 






Rennera 














X 






Oncosiphon 


o 












X 






Otospermum 










X 


X 








Heteromera 












X 








Daveaua 










X 


X 








Matricaria 


X 


X 


X 


X 


X 


X 


o 


o 





Microcephala 




X 


X 














Endopappus 












X 








Myxopappus 














X 






Foveolina 














X 






Lonas 










X 


X 








Tripleurospermum 


o 


X 


X 


X 


X 


X 


o 


o 


o 


Aaronsohnia 




X 








X 








Leucoptera 














X 






Adenoglossa 














X 






Hilliardia 














X 






Cotula 


X 


o 









X 


X 


X 


X 


Leptinella 
















X 


X 


Soliva 


X 


o 






o 


o 




X 


X 


Schistostephium 














X 






Hippia 














X 






Eriocephalus 














X 







The Matricariinae is our largest subtribe with respect to the 
number of genera (although the Artemisiinae, with Artemi- 
sia, contains more species). It consists mainly of a large group 
of predominantly annual and Mediterranean genera such as 
Matricaria and Tripleurospermum, and the predominantly 
South African Cotula group. Various South African genera 
are related to the Eurasian/North African taxa, for example, 
the annual genera Adenoglossa, Myxopappus , Foveolina, and 
possibly Rennera and Oncosiphon, as well as the South 
African shrubby genera Leucoptera, Cymbopappus, Pentzia 
and Marasmodes. 

The Cotula group is more narrowly and distinctly circum- 
scribed here than traditionally. It comprises the last seven 
genera in this account, characterized by their dorsiventrally 
flattened fruits. Hilliardia and Eriocephalus are both isolated 
and must be regarded as provisionally placed here, however. 

The whole subtribe is characterized by the arrangement of 
myxogenic cells on the cypselas, abaxially and on the ribs, but 
not on the adaxial surface. Furthermore, they have a pappus 
that is adaxially long, whether coroniform, auriculiform or of 
separate scales. Some genera do not have cypselas with 
myxogenic cells or with a pappus, for example, parts of the 
Cotula group. In the cladogram (Fig. 12) these two characters 
unite all Matricariinae, except the Cotula group, for reasons 
of parsimony. However, it is possible that they are synapo- 
morphies for the whole subtribe, since they occur also within 
the Cotula group. 

The positions of Oncosiphon and Rennera, both with 
epappose and non-myxogenic cypselas, are more doubtful. In 



groups in the cladogram. This is because they are similar in 
their mutual absence of some features, viz. myxogenic cells 
and pappus. In future studies they may turn out not to be 
closely related. The matter is further discussed by Kallersjo 
(1988). 

The generic interrelationships within the subtribe as shown 
in the cladogram (Fig. 12) must be regarded as very prelimi- 
nary. Many of the groupings are supported only by one or a 
few homoplasious characters, and numerous (226) equally 
parsimonious solutions are possible. We consider some sub- 
clades well supported, however. These include Ma3 with 
Cymbopappus , Pentzia and Marasmodes, Ma8 Heteromera 
and Daveaua, Ma9 Matricaria and Microcephala, Mal2 
Myxopappus and Foveolina, Ma 16 Leucoptera and Adeno- 
glossa, Ma 17 the Cotula group of genera (with the possible 
exclusion of Hilliardia and Eriocephalus), Ma 19 with Cotula, 
Leptinella, and Soliva, and Ma21 with Schistostephium, Hip- 
pia, and Eriocephalus (the last genus possibly to be 
excluded). All those clades are retained in the strict consen- 
sus tree of all the equally parsimonious cladograms found. 

Clades and characters - Fig. 12, Tables 2 and 25. 

Clade Mai - subtribe Matricariinae 

In this cladogram there are no synapomorphies for the 
subtribe but as explained above, characters 140 and 166 of 
clade Ma2 may be synapomorphies for the subtribe, with 
reversals in the Cotula group (Mal7). They appear as such in 
the alternative equally parsimonious cladograms. 



148 



K. BREMER AND C. J. HUMPHRIES 



— Ma3| 



|=Mal= 



F=Ma2:^ 



=Ma5=^ 



84 Cymbopappus 
ff= 85 Pentzia 

J"- 86 

rF=Ma6:ip 



86 Marasmodes 

87 Rennera 
Ma7=ii= 88 Oncosiphon 
89 Otospermum 

90 Heteromera 
l!=Ma8=!t= 91 Daveaua 
92 Matricaria 
L;=Ma9=!^ 93 Microcephala 

94 Endopappus 
?=Mall=|| rj= 95 Myxopappus 



=MalO 



r 

liMal2:!J= 96 Foveolina 
rj= 97 Lonas 
l!=Mal3=! !(= 98 Tripleurospermum 

J=Mal4| rp 99 Aaronsohnia 



FF- 102 Hilliardia 



100 Leucoptera 
Mal6=!J= 101 Adenoglossa 



'LMaieJC 



=Mal7i 



=Mal8 



rj=Mal9iJ=l 
iMa2ir 



103 Cotula 

rj= 104 Leptinella 



Ma2 0:t= 105 Soliva 
106 Schistostephium 
\\ r^ 107Hippia 

l!=Ma22ii= 108 Eriocephalus 



Fig. 12 Cladogram (of 226 possible) of the Matricariinae produced by the bb option in Hennig86. Cladogram length = 97, consistency index 
= 60, retention index = 68. 

Table 25 Data matrix for the Matricariinae. 1 = presence, = absence, ? = missing data or not applicable, p = polymorphic but scored as the 
plesiomorphic condition, a = polymorphic but scored as the apomorphic condition. 



1 
1 

5327 



11 



11 1 11 11111111 111 11111 111 1 1 11 111 

4505765 46 89333 77578226712174424372433784415541291279760736601545 
811 06220543132323387359816779059576531126824457564724063 711500 



1 111 

1755555666671 82 
3635679056806922 



84. Cymbopappus 

85. Pentzia 

86. Marasmodes 

87. Rennera 

88. Oncosiphon 

89. Otospermum 

90. Heteromera 

91. Daveaua 

92. Matricaria 

93. Microcephala 

94. Endopappus 

95. Myxopappus 

96. Foveolina 

97. Lonas 

98. Tripleurospermum 

99. Aaronsohnia 

100. Leucoptera 
10\. Adenoglossa 

102. Hilliardia 

103. Cotula 

104. Leptinella 

105. Soliva 

106. Schistostephium 

107. Hippia 

108. Eriocephalus 



111 
11? 
al? 
11? 
Ill 
111 
111 
111 
111 
111 
11a 
10? 
Ill 
11? 
Ill 
111 
111 
010 
111 
111 
111 
11? 
al? 
11? 
Ill 



?10 llall000000000000000000000000000?0000000000000000000000000? 

?lp Illlllp00pp0000000?00?000000p000?00000000000000000000?0000? 

?10 111111100000000000?00?0000000000?00000000000000000000?0000? 

?11 0?0001001111000000?00?0000000000?0000000000000G000000?0000? 

?11 0?010p0011101110000000p00000p00000000000000000000000000000? 

?11 110000001 1000001 1100000000000000?0000000000000000000000000? 

?11 110000011 1000000001 1000000000000?0000000000000000000000000? 

?11 11010001 11 00000000101 10000000000?0000000000000000000000000? 

Ill 1 100000a lOpOpOOOOOOOOplllOOpOOOOOOOOOOOOOOOOOOOOOOOpOOOOOO? 

?11 1000p0010000000010000alllOOOOOO?0000000000000000000000000? 

?11 110000001 0000000000 1000000 100000?011100000000000000p000000? 

?11 110001 00 lOOOpOOOOO? 10? 0000 100000?01001 11 0000000000000 ?00000 

?11 11000p001000p000000a000000100000?000011000000000000p000000? 

?11 11000 1011 000000000? 10?000010 11 1000000000000000000000000000? 

Ill al000p01a0p0000000010000000ap001110000000000000000000?0000? 

?11 11010p0110p000000000000000110000?000000000000000000p000000? 

?11 11 1000010000000000000000001 aOOOO?00100001 11 100000000000000? 

?11 1 1000001 100000000000000000100000?0010000111 11 1100000000000? 

?10 0? 10000000 101 00000000000001 00000??000000000a0001ap0app0000? 

?11 p?000p0ap010100000000000001p000001000000100a0001011app0000? 

?11 p?000p0a0010100000?00?00001p0000??000000a001000?0101 111000? 

?la 0?0000 1010 10 100000?00?0000100000??000000 1000000100000001 10? 

?10 0?100p000010100000?00?000010a000??00000010000000a00a000000? 

?10 0? 10000000 10000000?00?000010aOOO??000000 100100000 10 1000000? 

?10 0? 10000000 1000000000000000 10a 100? 1000000 100000000 lOpOOOOOl a 



??00000000000000 
???????????0ppp0 
???????????00000 
????????????0000 
?0000000000?0000 
??00000000000000 
??00000000000000 
??00000000000000 
?00000000000p000 
??00000000000000 
??00000000000000 
?0?????????00000 
?00000000000p000 
??00000000000000 
???????????00000 
??00000000000000 
??00000000000000 
??00000000000000 
?0000000000?00p0 
????????000?0000 
????????000?000p 
??000000000?0000 
????????000?0000 
????????000?0000 
??000000000?0000 



Clade Ma2 

140 Cypselas with myxogenic cells on abaxial surface and on 
the ribs of the adaxial surface. This character is assumed to be 
secondarily lost in Rennera and Oncosiphon. Several species 
of Tripleurospermum are also devoid of myxogenic cells, 
presumably secondarily. The character occurs also within 
Cotula and Leptinella. 

166 Pappus adaxially long. This character comprises a variety 



of different but homologous pappus types. The pappus is 
secondarily lost in Rennera and Oncosiphon. Some species of 
Matricaria, Tripleurospermum and Pentzia also lack pappus, 
presumably secondarily. 

Clade Ma3 

2 Plants shrubby. 

51 reversed. Floral parts without resin canals. 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



149 



82 Disc corolla tube thickened in fruit. This character is 
widespread in Achilleinae and Leucantheminae and occurs 
also in other genera of Matricariinae, notably Oncosiphon 
and Aaronsohnia. 

90 Disc corolla tube and and cypsela ribs with thick vascular 
strands. 

Cymbopappus 

There is no clear autapomorphy for this genus. 

Clade Ma4 

35 Capitula discoid. 

Pentzia 

There is no autapomorphy for this genus. 

Marasmodes 

34 Capitula sessile along the stems. 
Clade Ma5 

1 Plants annual. 

Clade Ma6 

73 Disc corolla lobes with dorsal appendages. 

Clade Ma7 

140 reversed. See clade Ma2. 

172 Pappus absent in ray and disc cypselas. 
Rennera 

35 Capitula discoid. 

153 Cypsela thick-walled and conspicuously rugose. 

Oncosiphon 

72 Disc corolla 4-lobed. Cotula and related genera also have 
4-lobed corollas. 

82 Disc corolla tube thickened in fruit. 

83 Disc corolla tube very thick and brittle. 

123 Cypselas with a mainly abaxial entire or toothed rim. 
Otospermum 

128 Cypselas with 1 abaxial and 2 lateral thick ribs and 2-3 
adaxial ribs. The arrangement of the three major ribs is 
abaxial-lateral and thus opposite to that of Tripleurospermum 
and many other genera. 

167 Pappus a stiff adaxial auricle. 

173 Testa epidermis cells spirally arranged around the embryo. 
A similar testa occurs in Microcephala. 

Clade Ma8 

115 Ray cypselas dorsiventrally flattened with 3 adaxial ribs. 

133 Cypselas with costal resin canals or sacs. 

Heteromera 

129 Cypselas with 1 adaxial and 2 lateral rather thick ribs. The 
same rib arrangement occurs in Tripleurospermum and 
related genera of clade MalO. 

Daveaua 



82 Disc corolla tube thickened in fruit. 

118 Ray cypselas laterally winged; wings projected to apical 
teeth. 

171 Pappus absent in disc cypselas, but present in ray cypselas. 

Clade Ma9 

46 Receptacle narrowly conical to subulate. 

47 Receptacle hollow. 

127 Cypselas with 5 mainly adaxially arranged ribs. 

Matricaria 

133 Cypselas with costal resin canals or sacs. 

Microcephala 

149 Cypselas with rather stiff unbranched hairs of 3-8 cells 
with spiral wall thickenings. 

173 Testa epidermis cells spirally arranged around the embryo. 
A similar testa occurs in Otospermum. 

Clade MalO 

129 Cypselas with 1 adaxial and 2 lateral rather thick ribs. The 
same rib arrangement occurs also in Heteromera. Foveolina 
has rather weak, but similarly arranged ribs. 

130 Cypselas with 2 lateral vascular strands, sometimes also 
with 1 adaxial strand. Tripleurospermum has a different 
autapomorphic type with 5 strands, with 2 abaxial strands 
associated with the two apical-abaxial resin sacs. 

Clade Mall 

141 Cypselas with dense rows of myxogenic cells also on the 
corona. This character is not expressed in Foveolina. 

Endopappus 

41 Involucral bracts wide, flabelliform. 

107 reversed. Cypselas not terete to weakly angled or flattened, 
but sharply angled. 

112 Cypselas large, with 3 thick protruding sclerenchymatous 
ribs, somewhat winged in ray cypselas. 

Clade Mal2 

156 Cypsela wall white and spongy; pericarpic cells isodiamet- 
ric with thin spiral wall thickenings. (Kallersjo, 1988). 

158 Cypsela wall with numerous druses in the pericarp. 
(Kallersjo, 1988). 

Myxopappus 

35 Capitula discoid. 

42 Involucral bracts subulate. 

43 reversed. Involucral bracts without scarious margins. 
Foveolina 

141 reversed. See clade Mall. 

Clade Mal3 

133 Cypselas with costal resin canals or sacs. 

Lonas 



150 



K. BREMER AND C. J. HUMPHRIES 



29 Capitula densely corymbose. 

35 Capitula discoid. 

45 Receptacle paleate. 

137 Cypselas with a single resin sac apically in the adaxial rib. 

Clade Mal4 

75 Disc corolla lobes with central resin sacs. The character is 
expressed in most but not all species of Tripleurospermum, in 
Aaronsohnia and in Leucoptera. It reverses in Adenoglossa. 

Tripleurospermum 

130 reversed. See clade MalO. 

136 Cypselas abaxially and apically with 2 distinct, occasion- 
ally 1 or 3-5, resin sacs. 

175 Embryo sac tetrasporic. 

183 Dehydrofalcarinone and dehydrofalcarinol present. These 
products in polyacetylene synthesis are common in the Aster- 
aceae but uncommon in Anthemideae, occurring also in 
Cotula and Eriocephalus. They may represent a secondarily 
simplified metabolism (Greger, 1977). The position of the 
character on the cladogram is uncertain because most genera 
of the subtribe have not been investigated chemically. 

Clade Mal5 

129 reversed. See clade MalO. 

Aaronsohnia 

82 Disc corolla tube thickened in fruit. 

Clade Mal6 

24 Leaves rather fleshy , few-lobed or entire. 
41 Involucral bracts wide, flabelliform. 

95 Anthers with endothecial tissue partly or wholly polarized. 

114 Cypselas dor siventr ally flattened. This is a traditional 
character of the Cotula group. 

117 Cypselas laterally winged. Winged fruits occur also within 
the Cotula group, clade Mal7. 

Leucoptera 

1 reversed. See clade Ma5. 

2 Plants shrubby. Several other genera in the Cotula group, as 
well as Pentzia and related genera, are also more or less 
shrubby. 

Adenoglossa 

15 reversed. Leaves not variously deeply lobed or divided, but 
entire. 

25 Leaves rather fleshy , entire, linear. 

52 reversed. Ray floret limb not white, but yellow. 

75 reversed. See clade Mal4. 

76 Disc corolla zygomorphic with 2 smaller adaxial lobes with 
marginal resin canals extending from the base of the corolla 
and with 3 larger abaxial lobes. 

94 Anthers with an apical resin sac. 



Clade Mal7 - The Cotula group of genera 

72 Disc corolla 4-lobed. Some Cotula and Soliva species have 
3-lobed corollas, another step in the reduction of the basically 
5-lobed corolla. Hippia and Eriocephalus have 5-lobed corol- 
las and the character is thus assumed to be reversed in these 
two genera. Oncosiphon also has 4-lobed corollas. 

130 Cypselas with 2 lateral vascular strands, sometimes also 
with 1 adaxial strand. 

172 Pappus absent in ray and disc cypselas. 

177 Chromosome number x=10. A variety of chromosome 
numbers are known from Cotula but 10 appears to be the 
base number. Most of the related genera are not investigated 
karyologically but Hilliardia also has x=10. Eriocephalus, 
however, has x=9, the plesiomorphic base number of the 
tribe. 

183 Dehydrofalcarinone and dehydrofalcarinol present. Most 
of the related genera have not been investigated chemically. 
However, these substances have been found also in Tripleu- 
rospermum. See further comments under that genus. 

Hilliardia 

2 Plants shrubby. 

51 reversed. Floral parts without resin canals. 

Ill Cypselas subglobose, with 2-3 very thin lateral-adaxial 
ribs. 

155 Cypsela wall very thin, translucent and showing brownish 
black, rounded, very thick-walled testa epidermis cells. 

Clade Mal8 

114 Cypselas dorsiventrally flattened. 

Clade Mal9 

117 Cypselas laterally winged. 

133 Cypselas with costal resin canals or sacs. 

Note: Leptinella and Soliva probably have their sister 
group(s) within Cotula as discussed under these genera. This 
is based on information other than that used to construct the 
cladogram. Thus Cotula could be paraphyletic and the char- 
acters appearing for Cotula in the cladogram should rather be 
interpreted as defining this clade, including all three genera, 
with reversals in Leptinella and Soliva. 

Cotula 

62 Outer florets stalked. This peculiar receptacular structure is 
a classical character for Cotula, though not particularly well 
developed in some species. Quite clearly it occurs also in 
Schistostephium and it is tempting to assume that it is a 
character for clade Mal8 although secondarily lost in the 
other genera of that clade. With this cladogram it is more 
parsimoniously interpreted as a parallelism between the two 
genera, however. 

Clade Ma20 

104 Disc floret style-branches fused. The same character 
occurs in Hippia and Eriocephalus. 

Leptinella 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



151 



70 Corolla inflated with a hollow space between outer surface 
and inner layer. 

Soliva 

1 Plants annual. Many species of Cotula are also annuals. 

34 Capitula sessile along the stems. 

36 Capitula disciform. Most species of Cotula as well as 
Hippia and Schistostephium are also disciform and it may be 
that the character is a synapomorphy for these genera. It is in 
conflict with other characters, however, and the presence of 
plesiomorphic, radiate species of Cotula, as well as the 
inclusion of Eriocephalus in clade Mal8, makes interpreta- 
tion difficult. 

63 Outer female florets in several rows. This character also 
occurs within Cotula, possibly indicating the sister group of 
Soliva within Cotula. See note under clade Mal9. 

67 Outer female florets without corollas. 

101 Style persistent and spinescent in fruit. 

133 reversed. See clade Mal9. 

Clade Ma21 

2 Plants shrubby. 

29 Capitula densely corymbose. 

51 reversed. Floral parts without resin canals. 

Ill reversed. See clade Mal7. 

Schistostephium 

36 Capitula disciform. See comment under Soliva. Some 
Schistostephium species are discoid, here considered a further 
development from the disciform condition. 

62 Outer florets stalked. See comment under Cotula. 

Clade Ma22 

72 reversed. See clade Mal7. 

104 Disc floret style-branches fused. 

Hippia 

36 Capitula disciform. See comment under Soliva. 

117 Cypselas laterally winged. 

Eriocephalus 

40 Involucral bracts in 2 unequal series. 

45 Receptacle paleate. 

50 Receptacular paleae pilose. 

84. CYMBOPAPPUS B. Nord. in Bot. Notiser 129: 
150 (1976). Type species: C. lasiopodus (Hutch.) B. 
Nord. (C. piliferus (Thell.) B. Nord.). 

Shrubs or half-shrubs. Leaves alternate, variously lobed or 
dentate, more or less cricoid. Capitula solitary, pedunculate, 
radiate. Receptacle convex to hemispherical or subcorneal, 
epaleate. Ray florets female, fertile; limb white or pinkish. 
Disc corolla 5-lobed; tube swollen and with thick vascular 
strands. Cypselas 5-ribbed, with large 2-celled glands and 
with myxogenic cells mainly abaxially; ribs with thick vascular 



strands. Pappus an oblique, adaxially longer whitish corona 
or cup, or of one large adaxial and one smaller abaxial scale. 

Distribution. S. Africa in the Cape, Transkei, and Trans- 
vaal. - 4 spp. 

Cymbopappus based on species formerly classified as Chry- 
santhemum and Marasmodes (Nordenstam, 1976) is close to 
Pentzia. They are distinguished by Cymbopappus being radi- 
ate and Pentzia discoid. Hence the species of Cymbopappus 
have not been usually associated with Pentzia although 
Nordenstam (1976) noted the relationship. Cymbopappus 
also has an almost tubular pappus but a similar pappus occurs 
in species of Pentzia. The distinction between these two 
genera deserves further study. 

C adenosolen (Harvey) B. Nord. 
C. hilliardiae B. Nord. 
C. piliferus (Thell.) B. Nord. 
C. lasiopodus (Hutch.) B. Nord. 

85. PENTZIA Thunb. in Prodr. pi. cap.: 145 (1800). 
Type species: P. crenata Thunb. {P. dentata (L.) 
OK.). 

Shrubs. Leaves alternate, variously pinnatisect, occasionally 
entire or apically dentate, more or less cricoid. Capitula 
sohtary or corymbose, pedunculate, discoid. Receptacle con- 
vex or hemispherical to conical, epaleate. Corolla 5-lobed; 
tube generally swollen and with thick vascular strands. 
Cypselas 5-ribbed, often with large 2-celled glands and with 
myxogenic cells abaxially and on the ribs; ribs with thick 
vascular strands. Pappus an adaxial auricle, or an oblique 
adaxially longer cup, or of free adaxially longer whitish 
scales, sometimes absent. 

Distribution. S. Africa mainly in the Cape, also in 
Namibia, two species {P. hesperidum and P. monodiana) also 
in N. Africa in Morocco and Algeria. - 23 spp. 

Traditionally (sensu Hutchinson, 1916fl,ft) Pentzia was all 
South African discoid Anthemideae with pinnatisect leaves 
and an epaleate receptacle. Two North African, typical 
Pentzia species, have also been described. Kallersjo (1988) 
has recently re-classified Pentzia and most South African 
species included in Tripleurospermum have been assigned to 
a number of genera, including Pentzia, Oncosiphon, Myxo- 
pappus and Foveolina. Pentzia now comprises discoid shrubs 
(no annuals) with more or less cricoid leaves and 5-ribbed, 
myxogenic cypselas. 

The separation of Pentzia from the small genera Cymbopa- 
ppus and Marasmodes is problematical. In future it might 
turn out that the latter genera have their sister group within a 
presently paraphyletic Pentzia. 

Clearly the generic delimitation of Pentzia is in need of 
detailed investigation, even after the removal of the annuals, 
as was undertaken by Kallersjo. 

At the species level Pentzia is also in need of revision. 
Some of the 'species' in the list below are probably conspe- 
cific but the list is taken from Kallersjo (1986). References 
are also given to species described after Hutchinson's 
{I9l6a,b) revision since they were not included in his key. 

P. argentea Hutch. 

P. bolusii Hutch. 

P. calcarea Kies - Note: Description in Kies, 1945. 



152 



K. BREMER AND C. J. HUMPHRIES 



P. calva S. Moore 

P. cooperi Harvey 

P. dentata (L.) OK. 

P. elegans DC. 

P. globosa Less. 

P. hesperidum Maire & Wilczek - Note: Description in 

Maire, 1936. 
P. incana (Thunb.) OK. 
P. lanata Hutch. 
P. monocephala S. Moore 
P. monodiana Maire & Wilczek - Note: Description in Maire, 

1929. 
P. nana Burch. 
P. peduncularis B. Nord. - Note: Description in Nordenstam, 

1987. 
P. pinnatisecta Hutch. 
P. punctata Harvey 
P. quinquefida (Thunb.) Less. 
P. sphaerocephala DC. 
P. spinescens Less. 
P. tomentosa B. Nord. - Note: Description in Nordenstam, 

1967. 
P. tortuosa (DC.) Fenzl ex Harvey 
P. viridis Kies - Note: Description in Kies, 1945. 

86. MARASMODES DC, Prodr. 6: 136 (1838). Type 
species: M. polycephalus DC. 

Shrubs. Leaves alternate, entire or occasionally lobed, linear, 
small, cricoid. Capitula small, closely aggregated at the 
stems, rarely solitary, discoid. Receptacle epaleate. Corolla 
5-lobed, glandular; tube swollen and with thick vascular 
strands. Anthers short and comparatively wide. Cypselas 
5-ribbed, glandular, with myxogenic cells mainly abaxially; 
ribs with thick vascular strands. Pappus of 7-10, oblong, flat, 
whitish, adaxially longer scales. 

Distribution. S. Africa in the Cape. - 4 spp. 

Marasmodes is a genus of cricoid shrubs characterized by 
small and few-flowered, aggregated and sessile capitula and a 
pappus of several, whitish scales. Pentzia generally has 
larger, pedunculate capitula and a coroniform/auriculate pap- 
pus, only rarely of a few separate scales. Otherwise Maras- 
modes is very similar to many species of Pentzia and it seems 
clear that Marasmodes has its sister group within Pentzia. In 
P. dentata the capitula are small and arranged in groups and 
the pappus is more or less cleft approaching the condition in 
Marasmodes. Adenosolen tenuifolius from eastern Cape and 
described by de Candolle (1837) was tentatively retained by 
Harvey (1865), although he considered it very similar in habit 
to his Marasmodes adenosolen (Cymbopappus adenosolen) 
from south-western Cape. The genus Adenosolen was 
reduced by later authors to a synonym of Marasmodes. 
However, it appears that Adenosolen tenuifolius is neither a 
Marasmodes nor a Cymbopappus , since according to the 
description it has cypselas without a pappus. The matter 
requires further investigation. 

Marasmodes was revised by Hutchinson (1917) but his key 
to the species cannot be recommended. The distinction 
between M. oligocephalus and M. polycephalus is better 
explained by Harvey (1865). M. adenosolen Harvey is Cym- 
bopappus adenosolen (Harvey) B. Nord. Hutchinson did not 
see the type, and erroneously accepted the identification of 



another collection (Schlechter 7899) as M. adenosolen. That 
collection represents M. oligocephalus. 

M. dummeri Bolus ex Hutch. 

M. oligocephalus DC. 

M. polycephalus DC. 

M. undulata Compton - Note: Description in Compton, 1946. 

87. RENNERA Merxm. in Mitt. bot. StSamml. 
Munch. 2: 335 (1957). Type species: R. limnophila 
Merxm. 

Annual herbs. Leaves alternate, pinnatisect. Capitula soli- 
tary, pedunculate, discoid. Receptacle hemispherical, epale- 
ate. Corolla 5-lobed, with a narrow tube and a wide enervate 
limb; lobes with dorsal appendages. Cypselas obovoid or 
5-angled, thick- walled and conspicuously tuberculate to rug- 
ose, apically sometimes with a thick, spreading to revolute 
rim. True pappus absent. 

Distribution. S. Africa in Namibia. - 3 spp. 

Rennera is characterized by the specialized, thick-walled 
cypselas. The thick apical rim, called a 'pappus' by 
Merxmiiller (1957), is present only in the type species. It does 
not seem to be homologous with the scarious pappus of 
Pentzia and many other Anthemideae. The thick spongy 
cypsela walls are presumably a dispersal adaptation; R. 
limnophila grows on temporarily submerged flats ('pans' and 
'vleis'). Two of the species were recently transferred to 
Rennera from Pentzia by Kallersjo (1988). 

The position of Rennera is unclear. It is not necessarily 
closely related to Pentzia (as circumscribed here), but provi- 
sionally placed in the same subtribe, Matricariinae. In the 
cladogram (Fig. 12) it is grouped with other annual genera 
from the Mediterranean and South Africa. The position as 
sister group to Oncosiphon is debatable, however. 

R. eenii (S. Moore) Kallersjo {Pentzia eenii S. Moore) 
R. limnophila Merxm. 

R. laxa (Bremek. & Oberm.) Kallersjo {Pentzia laxa 
Bremek. & Oberm.) 

88. ONCOSIPHON Kallersjo in Bot. J. Linn. Soc. 
96:310 (1988). Type species: O. piluliferum (L. f.) 
Kallersjo. 

Annual herbs. Leaves alternate, variously pinnatisect. 
Capitula solitary or corymbose, pedunculate, radiate or dis- 
coid. Receptacle flat to convex or conical to subglobose, 
epaleate. Ray florets female, fertile; limb white. Disc corolla 
4-lobed; with a very much swollen and brittle tube and a 
campanulate, partly enervate limb; lobes with dorsal append- 
ages. Cypselas 4-ribbed, without myxogenic cells, with a 
mainly abaxial entire or toothed rim. Pappus absent. 

Distribution. S. Africa mainly in the Cape, also in 
Namibia. - 8 spp. 

The species of this genus were formerly classified in Pentzia 
and Tripleurospermum. The position of the genus is difficult 
to assess. It may be related to other annual genera of subtribe 
Matricariinae, where it is provisionally placed. A relationship 
to some other part of Anthemideae is not evident. The 
swollen corolla tube and absence of pappus is characteristic 
also of subtribe Achilleinae, but Oncosiphon is different at 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 

least by its epaleate receptacle. The unusual chromosome 
numbers 2n=12 and 2n=16 (Nordenstam 1967, 1969; Mit- 
suoka & Ehrendorfer, 1972) are reported from species of this 
genus. 

O. africanum (P. Bergius) Kallersjo {Matricaria capensis L., 

Matricaria hirta (Thunb.) DC.) 
O. glabratum (Thunb.) Kallersjo 
O. grandiflorum (Thunb.) Kallersjo 
O. intermedium (Hutch.) Kallersjo 
O. piluliferum (L. f.) Kallersjo (Matricaria globifera (Thunb.) 

Fenzl ex Harvey, Pentzia globifera (Thunb.) Hutch.) 
O. sabulosum (Wolley-Dod) Kallersjo 
O. schlechteri (Bolus) Kallersjo 
O. suffruticosum (L.) Kallersjo {Matricaria multiflora Fenzl 

ex Harvey, Pentzia tanacetifolia (L.) Hutch.) 

89. OTOSPERMUM Willk. in Bot. Ztg 22: 251 
(1864). Type species: O. glabrum (Lagasca) Willk. 

An annual herb. Leaves alternate, pinnatisect. Capitula 
solitary or laxly corymbose, pedunculate, radiate. Receptacle 
conical, epaleate. Ray florets female, fertile; limb white; 
cypselas curved, basally connate to inner involucral bracts, 
with 1 abaxial and 2 lateral very thick ribs and 2-3 adaxial 
ribs, with myxogenic cells on the ribs; pappus an adaxial stiff 
auricle. Disc corolla 5-lobed; lobes with appendages; cypselas 
slightly curved, with 1 abaxial and 2 lateral thick ribs and 2 
adaxial ribs, with myxogenic cells on the ribs; testa of 
thick-walled, much elongated and spirally arranged cells; 
pappus an adaxial stiff auricle, shorter than in ray cypselas. 

Distribution. SW Europe in Spain and Portugal, N. Africa 
in Morocco, Algeria, and Tunisia. - Monotypic. 

The cypselas of Otospermum are unique in the Anthemideae. 
The three strong ribs are abaxially and laterally oriented, not 
adaxially and laterally as in Matricaria and some other 
genera. Nevertheless Otospermum seems best placed 
together with the other annual genera of Matricariinae. 

90. HETEROMERA Pomel, Nouv. mat. fl. atl.: 60 
(1874). Type species: H. fuscata (Desf.) Pomel. 

An annual herb. Leaves alternate, pinnatisect. Capitula 
solitary, rather long-pedunculate, radiate. Receptacle con- 
vex, epaleate. Ray florets female; limb white, many-veined; 
cypselas dorsiventrally flattened, with 3 adaxial ribs with 1-3 
resin canals; pappus a large adaxially longer scarious corona. 
Disc corolla 5-lobed; lobes with appendages; cypselas 
5-ribbed with 1 adaxial and 2 lateral stronger ribs and 2 
abaxial weaker ribs and with 3-5 resin sacs apically in the 
ribs, abaxially and on the ribs covered with myxogenic cells; 
pappus of 7-10 obovate, rounded, whitish, abaxially smaller 
scales. 

Distribution. N. Africa in Algeria, Tunisia and Libya. - 
Monotypic. 

Heteromera fuscata, often known as Chrysanthemum fusca- 
tum Desf., has sometimes been associated with two species 
known as Chrysanthemum deserticola (Murb.) F. Buxbaum 
and Chrysanthemum trifurcatum Desf. These have vallecular 
resin canals, however, and are here transferred to Chrysan- 
thoglossum (Leucantheminae). 

Heteromera fuscata has resin sacs in the ribs as in Aaron- 



153 

sohnia, Matricaria, Daveaua, and related genera. Daveaua 
appears to be the sister group. Both genera have the same 
kind of specialized ray cypselas, different from the disc 
cypselas. Alavi (1988) has used an eclectic treatment of the 
Matricariinae similar to that of Schultz Bipontinus (1860). In 
his treatment Heteromera is included in Tripleurospermum. 

H. philaenorum Maire & Weller, which is more pubescent 
than the type species and has connate pappus scales, is 
probably a synonym of H. fuscata (Jeffrey, \919b). 

91. DAVEAUA Willk. ex Mariz in Bolm Soc. 
broteriana 9: 243 (1891). Type species: D. 
anthemoides Mariz. 

An annual herb. Leaves alternate, pinnatisect. Capitula 
solitary, pedunculate, radiate. Receptacle conical, epaleate. 
Ray florets female; limb white; cypselas dorsiventrally flat- 
tened, with 3 adaxial ribs, laterally winged; wings projected 
to apical teeth; pappus a large adaxial auricle. Disc corolla 
5-lobed; tube basally swollen; lobes with appendages; cypse- 
las 5-ribbed with 3-5 resin sacs apically in the ribs and with 
myxogenic cells; pappus absent. 

Distribution. SW Europe in Portugal and N. Africa in 
Morocco. - Monotypic. 

Daveaua is apparently the sister group of Heteromera, both 
with the same kind of specialized ray cypselas. 

The shape of the cypsela resin sacs is interesting, being 
intermediate between the elongated canals of Aaronsohnia, 
Matricaria and Heteromera and the rounded sacs of Lonas 
and Tripleurospermum, indicating the homology of these 
structures. 

92. MATRICARIA L., Sp. pL: 891 (1753). Type 
species: M. recutita L. {Matricaria chamomilla L. 
(1755), non L. 1753 quae est M. inodora L. (1763), 
Chamomilla Gray). 

Annual herbs. Leaves alternate, pinnatisect. Capitula soli- 
tary, more or less pedunculate, radiate or disciform. Recep- 
tacle conical to subulate, hollow, epaleate. Ray florets 
female, fertile, with or without a white limb. Disc corolla 4- 
or 5-lobed; tube swollen in fruit; lobes rarely with resin 
canals. Cypselas shghtly dorsiventrally compressed, with 5 
mainly adaxially arranged thin ribs sometimes with resin 
canals, abaxially and on the ribs covered with myxogenic 
cells. Pappus absent or a small corona or sometimes, espe- 
cially in ray cypselas, an adaxial auricle. 

Distribution. N. hemisphere, widespread in Europe, 
Middle East, temperate Asia, N. Africa and N. America, 
some species (e.g. M. recutita) widespread as weeds also in 
the S. hemisphere. - 7 spp. 

Matricaria has characteristic, adaxially 5-ribbed fruits, and is 
thus clearly distinct from Tripleurospermum, with which it 
has been confused both taxonomically and nomenclaturally 
(see Pobedimova, 1961; Jeffrey, 1979^; Xifreda, 1985; 
Kergeulen et al., 1987). The question of typification was 
initially taken up by Hylander (1945) who considered that the 
name Matricaria chamomilla L. should be retained, although 
the description had been applied to the species now known as 
Tripleurospermum inodorum (L.) Schultz-Bip. in Linnaeus's 
Species plantarum (1753). Similarly, Toman & Stary (1965) 



154 



K. BREMER AND C. J. HUMPHRIES 



argued for the retention of Matricaria chamomilla L. on the 
grounds that it had been widely used in pharmalocogical 
literature. The selection of the Hortus Cliffortianus Linnean 
specimen of M. chamomilla L. as the lectotype was made by 
Grierson in Davis (1975) on the grounds that the phrase name 
'Matricaria foliis supra decompositis setaceis pedunculis soli- 
tariis' remained constant with the name Matricaria chamo- 
milla L. in all of Linnaeus's works. However, the Hortus 
Cliffortianus specimen of M. chamomilla has coronate cypse- 
las and does not therefore agree with Linnaeus's generic 
concept of Matricaria. The next choice of generic lectotypifi- 
cation, Matricaria recutita L., was made by Pobedimova, a 
species which agrees well with the generic description, 
because of its ecoronate cypselas (Jeffrey, 1979^) and there- 
fore is utilized here. 

The typification of Matricaria as based on Chamomilla 
vulgaris Gray was discussed by Rauschert (1974) who argued 
that Matricaria had been misapplied and must therefore be 
rejected. He argued that Chamomilla was the correct name 
for Matricaria and that Matricaria should be applied as the 
correct name for Tripleurospermum on the basis that M. 
chamomilla L. (1753) is equivalent to M. perforata Merat. He 
accordingly made the relevant new combinations in Tripleu- 
rospermum and Chamomilla. 

However, we reject Rauschert's argument since the typifi- 
cation of M. chamomilla L. (Grierson in Davis, 1975) and M. 
recutita L. are tied with the same generic concept. Indeed, 
Grierson considered that the two species were mere varieties 
of the same species. We accept Jeffrey's (1979fl) argument 
that because the type species was clearly designated by 
Pobedimova (1961) there is no confusion as to the correct 
application of Linnaeus's Matricaria and indeed the occa- 
sional misapplication to Tripleurospermum has not involved 
either type. Hence, M. recutita L. is the type species of 
Matricaria. 

Matricaria lasiocarpa Boiss. {Chamomilla lasiocarpa 
(Boiss.) Rauschert) is a species which we place in Micro- 
cephala. This genus has a number of autapomorphies not 
shared by Matricaria. Matricaria may be paraphyletic with 
Microcephala excluded. 

Tanacetum ledebourii (Schultz-Bip.) Schischk. from central 
Asia, also known as Matricaria songarica, is transferred into 
Matricaria. Tzvelev in Flora URSS (Komarov, 1961) included 
it in a monotypic, annual section of Cancrinia as C. discoidea. 
However, it has fruits typical of Matricaria and also agrees 
with Matricaria in several other characters. 

M. aurea (Loefl.) Schultz-Bip., (Chamomilla aurea (Loefl.) 
Gay ex Cosson & Krahk). Mediterranean, S. Europe, 
throughout N. Africa, the Middle East, and SW Asia to C. 
Asia. 

*M. macrotis Rech. f. (Chamomilla macrotis (Rech. f.) 
Rauschert). Turkey. 

M. matricarioides Porter ex Britton (M. discoidea DC., 
Chamomilla suaveolens (Pursh) Rydb.). W. North 
America, widespread as a cosmopolitan weed. 

M. occidentalis Greene (Chamomilla occidentalis (Greene) 
Rauschert). W. North America. 

M. recutita (L.) Rauschert (Matricaria chamomilla L. (1755) 
non L. 1753 (= Matricaria inodora L., Tripleurospermum 
inodorum (L.) Schultz-Bip.). Throughout Europe and 
most of temperate Asia, widespread as a cosmopolitan 
weed. 

M. songarica Bunge (Tanacetum ledebourii Schultz-Bip., 



Pyrethrum discoideum Ledeb.). C. Asia, Mongolia and 
China in Sinkiang. 
*M. tzvelevii Pobed. (Chamomilla tzvelevii (Pobed.) Raus- 
chert). Krym. 

93. MICROCEPHALA Pobed. in Bot. Mater. Gerb. 
bot. Inst. V. A. Komarova 21: 356 (1961). Type 
species: M. lamellata (Bunge) Pobed. 

Annual herbs. Leaves alternate, pinnatisect. Capitula soli- 
tary, rather long-pedunculate, radiate or discoid. Receptacle 
conical, hollow, epaleate. Ray florets female, fertile; limb 
white or pink. Disc corolla 5-lobed. Cypselas with 5 adaxially 
arranged ribs, abaxially and on the ribs with myxogenic cells, 
laterally with rather stiff, unbranched hairs of 3-8 cells with 
spiral cell wall thickenings; testa of much elongated spirally 
arranged cells. Pappus a large deeply fimbriate, whitish 
corona, in ray cypselas abaxially split to the base thus forming 
a large adaxial, deeply fimbriate auricle. 

Distribution. C. and middle Asia, Iran, Afghanistan Paki- 
stan. - 4 spp. 

Microcephala is well characterized by its peculiar cypsela 
hairs. The hair cells have spiral wall thickenings and are 
rather stiff. The five adaxially arranged cypsela ribs indicate a 
relationship with Matricaria. Apparently they are sister 
groups, or Microcephala may have its sister group within a 
paraphyletic Matricaria. In habit Microcephala is very similar 
to Matricaria. 

M. afghanica Podl. 

M. deserticola Podl. 

M. lamellata (Bunge) Pobed. (Chamomilla lasiocarpa 

(Boiss.) Rauschert, M. lasiocarpa (Boiss.) Pobed., M. 

turcomanica (Winkler) Pobed.). 
M. subglobosa (H. Kraschen.) Pobed. 

94. ENDOPAPPUS Schultz-Bip. in Bonplandia, 
Hannover 8: 369 (1860). Type species: E. 
macrocarpus Schultz-Bip. 

An annual herb. Leaves alternate, pinnatisect. Capitula 
soHtary, rather long-pedunculate, radiate. Involucral bracts 
with wide scarious margins. Receptacle flat, epaleate. Ray 
florets female, fertile; limb white or yellow, many-veined. 
Disc corolla 5-lobed. Cypselas comparatively large, with 1 
adaxial and 2 lateral thick sclerenchymatous and much pro- 
truding ribs, in ray cypselas somewhat winged, abaxially and 
on the ribs and pappus with myxogenic cells. Pappus a stiff 
short corona. 

Distribution. N. Africa in Morocco, Algeria and Tunisia. - 
Monotypic. 

Endopappus has no obvious close relatives but the cypsela 
shape with three adaxially and laterally thick ribs is similar to 
that of Tripleurospermum. Endopappus has no cypsela resin 
sacs, however. Nevertheless it is perhaps best considered 
together with Tripleurospermum. The South African Myxo- 
pappus and Foveolina are also possible relatives. 

The species is sometimes known as Chrysanthemum macro- 
carpum Cosson & Kralik or Pyrethrum macrocarpum (Cos- 
son & Kralik) Alavi. 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



155 



95. MYXOPAPPUS Kallersjo in Bot. J. Linn. Soc. 96: 
314 (1988). Type species: M. acutilobus (DC.) 
Kallersjo. 

Annual herbs. Leaves alternate, variously pinnatisect. 
Capitula solitary, pedunculate, discoid. Involucral bracts 
almost subulate, without scarious margins. Receptacle con- 
vex to conical, epaleate. Disc corolla 4- or 5-lobed. Cypselas 
with 1 adaxial and 2 lateral thick ribs, with myxogenic cells 
abaxially and on the ribs; cypsela wall white, spongy and 
containing numerous druses. Pappus an apical corona, cov- 
ered with myxogenic cells. 

Distribution. S. Africa in the Cape, Namibia. - 2 spp. 

The two species of Myxopappus were recently removed from 
Pentzia by Kallersjo. They seem related to Foveolina, 
another genus described by Kallersjo. Both Myxopappus and 
Foveolina have fruits similar to those of Tripleurospermum 
and Endopappus , and are hence grouped together with them 
in subtribe Matricariinae. The unusual chromosome number 
2n=14 was reported by Nordenstam (1967). 

M. acutilobus (DC.) Kallersjo 

M. hereroensis (O. Hoffm.) Kallersjo {Pentzia galpinii 
Hutch.) 



96. FOVEOLINA Kallersjo in Bot. J. Linn. Soc. 96: 
316 (1988). Type species: F. dichotoma (DC.) 
Kallersjo. 

Annual herbs. Leaves alternate, pinnatisect. Capitula soli- 
tary, pedunculate, radiate, disciform or discoid. Receptacle 
flat to convex to conical, epaleate. Ray florets female, fertile; 
limb white; cypselas adaxially somewhat flattened and incon- 
spicuously 3-ribbed, abaxially rounded and covered with 
myxogenic cells; cypsela wall adaxially white and spongy with 
numerous druses, abaxially very thin and transparent; pappus 
white and spongy, an adaxial auricle or occasionally coroni- 
form. If disciform, outer female florets filiform with cypselas 
almost terete, without myxogenic cells and without pappus; 
cypsela wall spongy and conspicuously wrinkled. Central 
florets with corolla 4- or 5-lobed; tube often basally dilated or 
saccate; cypselas either as in ray florets and with a pappus, or 
obovoid with a very thin cypsela wall all around and without a 
pappus, always abaxially with myxogenic cells. 

Distribution. S. Africa in the Cape, Namibia. - 5 spp. 

Foveolina comprises species formerly in Pentzia and 'Matri- 
caria'. The genus is related to Myxopappus, Tripleurosper- 
mum and other genera in subtribe Matricariinae. Two of the 
species, F. albidiformis and F. schinziana, are somewhat 
aberrant and resemble species of Cotula. The matter is 
discussed in detail by Kallersjo (1988). 

F. albida (DC.) Kallersjo {Pentzia annua DC.) 

F. albidiformis (Thell.) Kallersjo {Pentzia membranacea 

Hutch.) 
F. dichotoma (DC.) Kallersjo 
F. schinziana (Thell.) Kallersjo 
F. tenella (DC.) Kallersjo 



97. LONAS Adans., Fam. pi. 2: 118 (1763). Type 
species: Santolina annua L. (L. annua (L.) Vines & 
Druce). 

A glabrous annual herb. Leaves alternate, pinnatisect. 
Capitula in pedunculate dense corymbs, discoid. Receptacle 
elongated, paleate; paleae somewhat canaliculate, with a 
central resin canal. Corolla 5-lobed. Cypselas with 1 adaxial 
and 2 lateral vascular strands and ribs, with a single resin sac 
apically in the adaxial rib, abaxially and on the ribs covered 
with myxogenic cells. Pappus a scarious fimbriate corona. 

Distribution. Mediterranean, S. Europe in Italy and N. 
Africa in Morocco, Algeria, and Tunisia. - Monotypic. 

The position of Lonas is difficult to assess but we have placed 
it in the Matricariinae together with Tripleurospermum and 
related genera because of their similar fruits. Lonas differs by 
its densely corymbose capitula and by its paleate receptacle, 
both uncommon characters in Matricariinae. However, the 
arrangement of the myxogenic cells on the cypselas is typical 
of this subtribe. 

98. TRIPLEUROSPERMUM Schultz-Bip., 
Tanaceteen: 31-34 (1844fl). Type species: 
Tripleurospermum inodorum Schultz-Bip. 

Annual or perennial herbs. Leaves alternate, pinnatisect. 
Capitula solitary or corymbose, pedunculate, radiate, disci- 
form or discoid. Receptacle convex to conical, epaleate. Ray 
florets female; limb white or rarely pale pink. Disc corolla 
5-lobed; lobes each usually with a resin sac. Cypselas trique- 
trous with 1 adaxial and 2 lateral usually thick whitish ribs and 
sometimes 1-2 abaxial thin ribs, often rugose or tuberculate 
abaxially and between the ribs, sometimes with myxogenic 
cells, abaxially and apically with 2 distinct, occasionally 1 or 
3-5, resin sacs. Pappus a corona or auricle, or of few scales, 
sometimes absent. Embryo sac tetrasporic. Flavonol 
7-glycosides and dehydrofalcarinone and dehydrofalcarinol 
present. 

Distribution. Europe and temperate Asia, a few species 
also in N. America {T. maritimum and T. perforatum natural- 
ized) and N. Africa {T. auriculatum) , one species {T. perfora- 
tum) widespread as a weed; most species in SE Europe and 
SW Asia. - 38 spp. 

Despite the comparatively large number of species, Tripleu- 
rospermum as presently circumscribed is a distinct genus with 
characteristic 3-ribbed cypselas with 2 abaxial-apical resin 
sacs. Chemically about 30 species have been investigated for 
flavonol glycosides and they all contain 7-glycosides rather 
than the common 3-glycosides as, for example, in Matricaria 
(Greger, 1975, 1977). Species of Tripleurospermum embryo- 
logically investigated have a tetrasporic embryo sac. 

In habit Tripleurospermum is similar to many other 
Anthemideae. The genus was earUer more widely and less 
distinctly circumscribed, so as to also include Matricaria. 
However, we keep the genus distinct following Pobedimova 
in Flora USSR (Komarov, 1961) and Grierson in Flora of 
Turkey (Davis, 1975). Tripleurospermum as here understood 
follows the original concept of Schultz Bipontinus (1844^). 
The South African species described under Matricaria are 
now transferred to other genera (Kallersjo, 1988). 

As described under Matricaria Rauschert (1974) and Kay 
(1976) missapplied the name Matricaria to refer exclusively to 



156 



K. BREMER AND C. J. HUMPHRIES 



the species of Tripleurospermum. Because Rauschert (1974) 
transferred a number of species from Tripleurospermum to 
Matricaria we have cited a more complete synonomy. The 
species list is compiled from the standard floras with later 
species and species described from other areas added, for 
example E. Hossain in Flora of Turkey (Davis, 1975). 

T. auriculatum (Boiss.) Rech. fil. {Matricaria auriculata 
(Boiss.) Ball). Middle East. 

T. baytopianum E. Hossain. Turkey. 

T. breviradiatum (Ledeb.) Pobed. {Matricaria breviradiata 
(Ledeb.) Rauschert). W. Siberia. 

T. callosum (Boiss. & Heldr.) E. Hossain {Chamaemelum 
callosum Boiss. & Heldr.). Turkey. 

T. caucasicum (Willd.) Hayek {Matricaria caucasica (Willd.) 
Poiret). SE Europe, Turkey, Caucasus, Middle East, Iraq, 
Iran, Afghanistan. 

T. colchicum (Manden.) Pobed. {Matricaria colchica 
(Manden.) Rauschert). Caucasus. 

*T. conoclinium (Boiss. & Balansa) Hayek {Matricaria cono- 
clinia (Boiss. & Balansa) Nyman). Turkey. 

*T. corymbosum E. Hossain (non Matricaria corymbosa 
Desr. in Lam.). Turkey. 

*T. daghestanicum (Rupr. ex Boiss.) Bremer &. Humphries, 
comb. nov. Basionym: Chamaemelum daghestanicum 
Rupr. ex Boiss., Fl. orient. 3: 334 (1875) {Matricaria 
daghestanica (Rupr. ex Boiss.) Rauschert). 

T. decipiens (Fischer & C. Meyer) Bornm. {Matricaria decipi- 
ens (Fischer & C. Meyer) K. Koch ). Turkey, Caucasus, 
Iran. 

T. disciforme (C. Meyer) Schultz-Bip. {Matricaria disciformis 
(C. Meyer) DC. ). Turkey, Caucasus, Iraq, Iran, Afghani- 
stan, Pakistan, C. Asia. 

*T. elongatum (Fischer & C. Meyer ex DC.) Bornm. {Matri- 
caria elongata (Fischer & C. Meyer ex DC.) Hand.-Mazz., 
M. australis (Pobed.) Rauschert, Tripleurospermum aus- 
trale Pobed.). Turkey, Caucasus. - Note: T. australe was 
included as a doubtful synonym by E. Hossain in Flora of 
Turkey (Davis, 1975). 

*T. fissurale (Sosn.) E. Hossain. Turkey. 

*T. froedinii Rech. f. {Matricaria froedinii (Rech. f.) Raus- 
chert). Iran. 

*T. grossheimii (Fed.) Pobed. {Matricaria grossheimii (Fed.) 
Rauschert). Caucasus. 

T. heterolepis (Freyn & Sint.) Bornm. {Chamaemelum het- 
erolepis Freyn & Sint.). Turkey. 

*T. homogamum G. X. Fu. China. 

T. hygrophilum (Bornm.) Bornm. {Matricaria hygrophila 
(Bornm.) Rauschert). Turkey. 

T. karjaginii (Manden. & Sof.) Pobed. {Matricaria karjaginii 
(Manden. & Sof.) Rauschert). Caucasus. 

T. kotschyi (Boiss.) E. Hossain {Chamaemelum kotschyi 
Boiss.). Turkey. 

T. limosum (Maxim.) Pobed. {Matricaria limosa (Maxim.) 
Kudo). Far East, Japan, China. 

T. maritimum (L.) K. Koch {Matricaria maritimum L. Tri- 
pleurospermum ambiguum (Ledeb.) Franchet & Savat., T. 
phaeocephalum (Rupr.) Pobed., T. subpolare Pobed.). 
Throughout most of Europe, temperate Asia and in North 
America. - Note: in Flora europaea three subspecies are 
recognized: ssp. maritimum, ssp. phaeocephalum (Rupr.) 
Rauschert, and ssp. subpolare (Pobed.) Rauschert. The 
related weedy species T. perforatum {= T. inodorum) has 
been kept separate. 



T. microcephalum (Boiss.) Bornm. (non Matricaria micro- 
cephala C. Koch, M. armeniaca Rauschert). Turkey, Iran, 
Iraq. 

T. monticolum (Boiss. & Huet) Bornm. {Matricaria monti- 
cola (Boiss. & Huet) Rauschert). Turkey. 

T. oreades (Boiss.) Rech f. {Matricaria oreades Boiss., Matri- 
caria halepensis Rauschert, Matricaria szowitzii (DC.) 
Rauschert, Tripleurospermum szowitzii (DC.) Pobed., 
Matricaria tchihatchewii (Boiss.) Voss, Tripleurospermum 
tchihatchewii (Boiss.) Bornm.). Turkey, Middle East, Cau- 
casus, Iran. - Note: Matricaria halepensis was a new name 
for Chamaemelum grandiflorum Boiss. & Hausskn., the 
epithet of which is already occupied in Matricaria. How- 
ever, this species as well as the others listed in the 
synonymy above were reduced by E. Hossain in Flora of 
Turkey (Davis, 1975). 

T. parviflorum (Willd.) Podeb. {Matricaria parviflora 
(Willd.) Poiret). E. Europe in Russia, Turkey, Caucasus, 
Middle East, Iraq, Iran, Pakistan, C. Asia. 

T. perforatum (Merat) Lainz {Matricaria perforata Merat, 
Matricaria inodora L., Tripleurospermum inodorum (L.) 
Schultz-Bip.). Throughout most of Europe and temperate 
Asia, widespread as a weed especially in North America 
etc. - Note: See T. maritimum. 

*T. pichleri (Boiss.) Bornm. {Matricaria pichleri (Boiss.) 
Rauschert). Turkey. 

T. repens (Freyn & Sint.) Bornm. Turkey. 

T. rosellum (Boiss. & Orph.) Hayek {Matricaria rosella 
(Boiss. & Orph.) Nyman, Matricaria lesbiaca (Candargy) 
Rauschert, Tripleurospermum lesbiacum (Candargy) Rech. 
f.). SE Europe in Greece, Turkey. - Note: The synony- 
mous names are from E. Hossain in Flora of Turkey 
(Davis, 1975). 

*T. rupestre (Sommier & Levier) Pobed. {Matricaria rupestris 
(Sommier & Levier) Rauschert). Caucasus. 

T. sevanense (Manden.) Pobed. {Matricaria sevanensis 
(Manden.) Rauschert). Turkey, Caucasus, Iran. 

T. subnivale Pobed. {Matricaria subnivalis (Pobed.) Raus- 
chert). Caucasus. 

T. tempskyanum (Freyn & Sint.) Hayek {Matricaria 
tempskyana (Freyn & Sint.) Rauschert). SE Europe in 
Greece. 

T. tenuifolium (Kit.) Freyn {Matricaria trichophylla (Boiss.) 
Boiss.). SE and C. Europe, Turkey. 

T. tetragonospermum (Schum.) Pobed. {Matricaria tetragono- 
sperma (Schum.) Hara & Kitam.). W. Siberia, Far East, 
Japan, China. 

T. transcaucasicum (Manden.) Pobed. {Matricaria transcau- 
casica (Manden.) Rauschert). Turkey, Caucasus. 

*T. tzvelevii Pobed. (non Matricaria tzvelevii Pobed.), {M. 
aserbaidshanica Rauschert). Caucasus. 

99. AARONSOHNIA Warb. & Eig in Bull, agric. 
Exp. Stn, Tel-Aviv 6: 39 (1927). Type species: A. 
factorovskyi Warb. & Eig. 

Annual herbs. Leaves alternate, pinnatisect. Capitula soli- 
tary, rather long-pedunculate, radiate, disciform, or discoid. 
Receptacle conical, epaleate. Ray florets female, fertile, with 
or without a white limb. Disc corolla 5-lobed; tube basally 
and especially abaxially much swollen in fruit; lobes with 
central resin canals. Cypselas slightly dorsiventrally flattened 
with 1 adaxial and 2 lateral vascular strands and often with 2 
lateral resin canals, abaxially covered with myxogenic cells. 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



157 



Pappus an adaxial, conspicuous, whitish auricle, sometimes 
absent. 

Distribution. Middle East and N. Africa in Morocco, 
Algeria, Tunisia and Libya. - 2 spp. 

During this study it became clear that Chlamydophora pube- 
scens in almost all details agrees with the circumscription of 
the monotypic genus Aaronsohnia from the Middle East. 
Thus, we have transferred it to Aaronsohnia. A. pubescens is 
a sometimes radiate or discoid species from North Africa. 

Aaronsohnia is similar to Cotula and other herbaceous 
members of the Cotula group. However, it is probably more 
closely related to Matricaria, Leucoptera and Adenoglossa as 
indicated by the character analysis. 

A. factorovskyi Warb. & Eig 

A. pubescens (Desf.) Bremer & Humphries, comb. nov. 
Basionym: Cotula pubescens Desf., Fl. atlant. 2: 284 (1799) 
{Chlamydophora pubescens (Desf.) Cosson & Durieu, 
Matricaria pubescens (Desf.) Schultz-Bip., Chamomilla 
pubescens (Desf.) Alavi). 

100. LEUCOPTERA B. Nord. in Bot. Notiser 129: 
141 (1976). Type species: L. nodosa (Thunb.) B. 
Nord. 

Shrublets. Leaves alternate or opposite, entire or few-lobed, 
somewhat fleshy. Capitula solitary, long-pedunculate, radi- 
ate. Involucral bracts wide, many-veined. Receptacle convex, 
epaleate. Ray florets female, fertile; Hmb white, often 
becoming pink-reddish. Disc corolla 5-lobed; lobes generally 
with central resin sacs. Anthers with endothecial tissue 
mainly polarized. Cypselas dorsiventrally flattened, laterally 
winged, whitish, with 1 adaxial and 2 lateral vascular strands, 
generally with 2 lateral resin canals, abaxially covered with 
elongated apically projected myxogenic cells. Pappus of 1 
larger adaxial auricle and 2 smaller adaxial-lateral scales. 

Distribution. S. Africa in the Cape. - 3 spp. 

This distinct, recently described genus of three species is 
closely related to Adenoglossa. Indeed, they are sister groups 
as stated by Nordenstam (1976). He also considered Leucop- 
tera monophyletic, despite the fact that it is plesiomorphic in 
most characters in comparison to Adenoglossa. 

Leucoptera and Adenoglossa are similar to the Cotula 
group in fruit morphology. However, they are possibly more 
closely related to the northern hemisphere species of Matri- 
caria and Aaronsohnia as indicated by the cladogram (Fig. 
12). Their position within the Matricariinae is uncertain. 

L. nodosa (Thunb.) B. Nord. 
L. oppositifolia B. Nord. 
L. subcarnosa B. Nord. 

101. ADENOGLOSSA B. Nord. in Bot. Notiser 129: 
137 (1976). Type species: A. decurrens (Hutch.) B. 
Nord. 

An annual herb. Leaves alternate or opposite, entire, fleshy. 
Capitula solitary, long-pedunculate, radiate. Involucral 
bracts wide, many-veined. Receptacle conical, epaleate. Ray 
florets female, fertile; limb short, yellow. Disc corolla 
5-lobed, slightly zygomorphic with 2 smaller adaxial lobes 
with marginal resin canals extending from the base of the 



corolla and with 3 larger abaxial lobes. Anthers with an apical 
resin sac; endothecial tissue partly polarized. Cypselas dor- 
siventrally flattened, laterally winged, whitish, with 1 adaxial 
and 2 lateral vascular strands and with 2 lateral resin canals, 
abaxially covered with elongated apically projected myxo- 
genic cells. Pappus of 1 larger adaxial auricle, 2 larger 
adaxial-lateral scales, and 2-3 smaller abaxial scales. 

Distribution. S. Africa in the NW Cape. - Monotypic. 

Adenoglossa is a recently described monotypic genus with a 
number of autapomorphies. It is closely related to Leucoptera 
and their interrelationship is discussed under that genus. 

102. HILLIARDIA B. Nord. in Op. bot. Soc. bot. 
Lund 91'. 147 (1987). Type species: H. zuurbergensis 
(Oliver) B. Nord. 

A scrambling shrub. Leaves alternate, flat, pinnatisect. 
Capitula laxly corymbose, pedunculate, radiate. Receptacle 
conical to elongated, epaleate. Ray florets female, fertile; 
limb white, many-veined, apically rather deeply bifid. Disc 
corolla 4-lobed. Cypselas subglobose, with 2-3 very thin 
adaxial ribs; cypsela wall very thin, translucent and showing 
the brownish black, rounded and very thick-walled testa 
epidermis cells. Pappus absent. 



Distribution. S. Africa in the Cape and Natal, 
typic. 



Mono- 



Hilliardia was formerly included in Matricaria but it is related 
neither to the northern hemisphere Matricaria s. s. nor to the 
South African Matricaria species transferred to various other 
genera (Myxopappus, Foveolina, Oncosiphon, Cotula). Hill- 
iardia has unusual thin-walled fruits, with an extremely well 
developed testa (seen in mature fruits only). The shape and 
venation of the fruits recall Achillea and Cotula. Hilliardia is 
also similar to Cotula and its South African relatives have a 
4-lobed corolla and a chromosome number of x=10, hence 
our placement of Hilliardia within the Cotula group. 

103. COTULA L., Sp. pL: 891 (1753). Type species: 
C. coronopifolia L. - Brocchia Vis., Cenia Comm. 
ex Juss., Otochlamys DC, Sphaerodinium (DC.) 
Schultz-Bip. 

Annual or perennial herbs. Leaves alternate or sometimes 
opposite or rosulate, pinnatisect, lobed or occasionally 
entire. Capitula solitary, pedunculate, generally disciform or 
discoid or rarely shortly radiate; peduncles sometimes 
inflated below the capitulum. Receptacle flat to conical, 
epaleate. Ray or outer female florets fertile, in one to several 
rows, generally stalked; tube short or absent; limb generally 
absent, white if present. Disc corolla 4-lobed, occasionally 
3-lobed, abaxially sometimes saccate; lobes generally with 
central resin canals, one lobe occasionally expanded to a 
radiate limb. Cypselas dorsiventrally flattened, often laterally 
winged, with 2 lateral (occasionally 3 or 4) vascular strands 
and sometimes with 2 lateral resin canals, generally with 
blunt uniseriate hairs, sometimes with myxogenic cells. Pap- 
pus absent. Flavonol 5-glycosides and dehydrofalcarinone 
and dehydrofalcarinol present. 

Distribution. S. hemisphere, mainly in S. Africa, also in 
Australia, S. America (C. mexicana), and on the S. oceanic 
islands (C. goughensis, C. moseleyi), a few species extending 



158 



K. BREMER AND C. J. HUMPHRIES 



to E. Africa (C. abyssinica, C. cryptocephala), N. Africa (C. 
cinerea), and to Mexico (C. mexicana); a few widespread 
weedy species (C. anthemoides , C. aus trails, C. coronopifo- 
lia). - 55 spp. 

Cotula is generally conceived as disciform or discoid, except 
for some South African species with rays secondarily evolved 
from one of the disc floret corolla lobes. But there is also a 
South African C. montana (Adamson et al., 1944) with true 
rays. It is clearly a Cotula with its typical fruits and stalked 
rays. Thus, it appears that Cotula may be radiate as well as 
disciform or discoid. 

The monotypic genus Sphaeroclinium, revived by Mit- 
suoka & Ehrendorfer (1972), is included as a synonym of 
Cotula. The South African species, 5. nigellifolium, was 
originally described as a species of Matricaria, a 'dustbin' 
genus in South Africa used for various species of unknown 
relationships. Since the species is radiate its close relationship 
to the generally disciform species of Cotula is not immedi- 
ately obvious. Furthermore, the marginal florets, i. e. the 
rays, are not stalked in Sphaeroclinium as in most species of 
Cotula. However, there are several species of Cotula with 
very shortly stalked or sessile rays and there is at least one 
described radiate species C. montana. Thus, the distinction 
between Sphaeroclinium and Cotula disappears on closer 
examination. Both genera have a similar habit, 4-lobed 
corollas, the same specialized cypsela morphology, a chromo- 
some number of x=10, and related chemistry. 

Another South African species of 'Matricaria' belonging 
here is M. andreae E. Phillips. It was described as a Matri- 
caria presumably because it is radiate but the fruit morphol- 
ogy leaves no doubt about its generic position. Thus, it is 
recombined here as Cotula andreae. 

Lloyd (1972a, ft) recognized three sections within Cotula: 
sect. Cotula with usually one row of rays, sect. Strongy- 
losperma with several rows of rays and sect. Leptinella with 
inflated corollas. Sect. Leptinella is recognized as a distinct 
genus (Lloyd & Webb, 1987) but the demarcation between 
the other two sections is difficult since they can hardly be 
defined as monophyletic groups. The South African segre- 
gate genera Cenia, with inflated peduncles, and Otochlamys, 
with an abaxially saccate corolla, are included in sect. Cotula. 
Most species belong to this section, which is mainly South 
African. Sect. Cotula together with sect. Strongylosperma are 
clearly in need of revision and the sectional delimitation 
should be reconsidered. 

It is also possible that Cotula is paraphyletic with Leptinella 
and Soliva excluded. Saliva with several rows of ray florets is 
possibly derived from sect. Strongylosperma or part of it. In 
the cladogram Leptinella and Soliva are sister genera, united 
by their functionally male disc florets, but interrelationships 
between these two genera and the subunits of Cotula deserve 
further investigation. 

Identification of Cotula species, especially South African, 
is difficult, since the genus is not revised and since the 
descriptions are scattered in the literature. In South Africa as 
a whole only Flora capensis (Harvey, 1865) is available and 
there are numerous species described later. References are 
given in the list of species. However, for Natal with its eight 
species there is a modern treatment by Hilliard (1977). 

C. abyssinica Schultz-Bip. E. Africa. 

C. alpina (Hook, f.) Hook. f. Australia. 

C. andreae (E. Phillips) Bremer & Humphries, comb. nov. 



Basionym: Matricaria andreae E. Phillips in//. S. Afr. Bot. 
16: 21 (1950). S. Africa. 

C. anthemoides L. S. Africa, widespread as a cosmopolitan 
weed. 

C. australis (Sieber ex Sprengel) Hook. f. Australia, wide- 
spread as a cosmopolitan weed. 

C. barbata DC. S. Africa. 

C. bipinnata Thunb. S. Africa, naturalized in Australia. 

*C. bracteolata E. Meyer ex DC. S. Africa. 

C. ceniifolia DC. S. Africa. 

C. cinerea Del. {Brocchia cinerea (Del.) Vis.). N. Africa. - 
Note: This species seems aberrant within Cotula. It has 
been described as the genus Brocchia, which possibly 
should be re-established. The matter requires further 
study. 

C. coronopifolia L. S. Africa, Australia, widespread as a 
cosmopolitan weed. 

*C. cotuloides (Steetz) Druce. Australia. 

C. cryptocephala Schultz-Bip. ex A. Richard. E. Africa. 

*C. dielsii Muschler. S. Africa. 

C. duckittiae (L. Bolus) Bremer & Humphries, comb. nov. 
Basionym: Cenia duckittiae L. Bolus in Ann. Bolus Herb. 
4: 15 (1925) (Cenia expansa Compton). S. Africa. - Note: 
C. expansa is provisionally included here as a synonym. We 
find it hardly distinct and hesitate to make a new combina- 
tion unnecessarily. 

C. eckloniana (DC.) Levyns {Otochlamys eckloniana DC). 
S. Africa. 

*C. elongata Vogel. Java. 

C. filifolia Thunb. S. Africa. 

*C. goughensis R. N. R. Brown. Gough Island. 

C. heterocarpa DC. S. Africa. 

C. hispida (DC.) Harvey. S. Africa. 

C. laxa DC. S. Africa. 

C. leptalea DC. S. Africa. 

C. lineariloba (DC.) Hilliard. S. Africa. 

*C. loganii Hutch. S. Africa. 

*C. macroglossa Bolus ex Schltr. S. Africa. 

C. mariae Bremer & Humphries, nom. nov. Basionym: Cenia 
pectinata DC, Prodr. 6: 83 (1837) (non Cotula pectinata 
Hook, f.) - Note: We name this species after Ms. Mari 
Kallersjo in recognition of her contributions to the classifi- 
cation of South African Anthemideae. S. Africa. 

C. melaleuca Bolus. S. Africa. 

C. membranifolia Hilliard. S. Africa. 

C. mexicana (DC) Cabrera (C. pygmaea Benth. in Benth. & 
Hook, f., C. pedicellata (Ruiz Lopez & Pavon) Cabrera 
(non Compton), C. cabrerae Caro). S. and C America, 
Mexico. 

C. microglossa (DC) O. Hoffm. & Kunze ex Kunze. {Cenia 
albovillosa S. Moore, Cenia microglossa DC.) - Note: C. 
albovillosa is provisionally included as a synonym. We find 
it hardly distinct and hesitate to make a new combination 
unnecessarily. S. Africa. 

C. montana Compton. S. Africa. 

C. moseleyi Hemsley. Tristan d'Acuna. 

C. myriophylloides Harvey in Hook. S. Africa. 

C. nigellifolia (DC) Bremer & Humphries, comb. nov. 
Basionym: Matricaria nigellifolia DC, Prodr. 6: 50 (1837) 
{Sphaeroclinium nigellifolium (DC) Schultz-Bip.). S. 
Africa. 

C. nudicaulis Thunb. S. Africa. 

C. paludosa Hilliard. S. Africa. 

*C. paradoxa Schinz. S. Africa. 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



159 



*C. pedicellata Compton. S. Africa. 

C. pedunculata (Schltr) E. Phillips {Otochlamys pedunculata 

Schltr. S. Africa. 
*C. pterocarpa DC. S. Africa. 
C pusilla Thunb. S. Africa. 
*C. radiata O. Hoffm. ex OK. S. Africa. 
C. radicalis (Killick & Claassen) Hilliard & Burtt. S. Africa. 
*C. rosea Boj. ex Less. Madagascar. 
C. sericea L. f. (Cenia sericea (L. f.) DC). S. Africa. 
C. socialis Hilliard. S. Africa. 
C. sororia DC. S. Africa. 
*C. stenophylla K. Koch. S. Africa. 
C tenella E. Meyer ex DC. S. Africa. 
C. thunbergii Harvey. S. Africa. 
C. turbinata L. {Cenia turbinata (L.) Pers.). S. Africa, 

naturalized in Australia. 
C. villosa DC. (C. multifida DC). S. Africa. 
C. vulgaris Levyns. S. Africa, Australia. 
C. zeyheri Fenzl ex Harvey. S. Africa. 



104. LEPTINELLA Cass, in Bull. Sci. Soc. philom. 
Paris 1822: 127 (1822). Type species: L. scariosa 
(Cass.) Franchet (Lloyd, 1972fl). 

Perennial or facultative annual herbs, prostrate or rarely 
suberect. Leaves alternate or opposite, pinnatisect. Capitula 
soHtary, pedunculate, axillary, discoid or disciform, with 
varying proportions of pistillate and staminate florets or 
unisexual or both. Receptacle flat or conical, epaleate. 
Outer, female florets fertile, in one to several rows, tubular, 
conical to cylindrical, inflated with a hollow space between 
the outer surface and an inner layer closely surrounding the 
style. Central florets female-sterile, functionally staminate; 
corolla 4-lobed. Cypselas dorsiventrally flattened or more 
often thick and convex on the dorsal surface at least, glabrous 
or rarely with tapering uniseriate hairs on the margins. 
Pappus absent. 

Distribution. New Guinea, Australia, New Zealand and its 
subantarctic islands and one species in South America and 
the Falkland Islands. - 33 spp. 

Until recently resurrected by Lloyd & Webb (1987), Lepti- 
nella has been included in the genus Cotula since being 
relegated to it as an infrageneric section by Hooker (1864). 
Lloyd (1972a, b) divided Cotula into three sections: sect. 
Cotula, sect. Strongylosperma (Less.) Benth. and sect. Lepti- 
nella (Cass.) Benth. Although Lloyd & Webb (1987) consider 
that it is very likely that the three sections form a monophyl- 
etic group we think so only with the addition of the genus 
Soliva. As described below Soliva is related to sect. Strongy- 
losperma and these two groups together with Cotula and 
Leptinella are most likely a monophyletic group. Lloyd & 
Webb (1987) argue that because Leptinella is defined by the 
three distinctive autapomorphies it should be recognized as a 
genus in its own right. All species of Leptinella share the same 
habit, the same peculiar female floret morphology and all 
species examined so far have a chromosome number of x=13. 
Furthermore, the distribution is clearly West Pacific occur- 
ring in New Guinea, southeast Australia, New Zealand and 
the subantarctic islands. For the New Zealand species and the 
New Guinea species the treatments by Lloyd (1972fl) and van 
Royen & Lloyd (1975), respectively, are available. 



L. albida (D. Lloyd) D. Lloyd & C Webb {Cotula sericea 

(Kirk) Cockayne & Allan). New Zealand. 
*L. altilitoralis (P. Royen & D. Lloyd) D. Lloyd & C Webb. 

New Guinea. 
L. atrata (Hook, f.) D. Lloyd & C Webb. New Zealand. 
L. calcarea (D. Lloyd) D. Lloyd & C Webb. New Zealand. 
L. dendyi (Cockayne) D. Lloyd & C Webb. New Zealand. 
L. dioica Hook. f. New Zealand. 

L. dispersa (D. Lloyd) D. Lloyd & C Webb. New Zealand. 
*L. drummondii (Benth.) D. Lloyd & C Webb. Australia. 
L. featherstonii F. Muell. {Cotula renwickii Cockayne). New 

Zealand. 
*L. filicula (Hook, f.) Hook. f. Australia (Tasmania). 
L. filiformis (Hook, f.) D. Lloyd & C Webb. New Zealand. 
L. goyenii (Petrie) D. Lloyd & C Webb. New Zealand. 
L. intermedia (D. Lloyd) D. Lloyd & C Webb. New 

Zealand. 
L. lanata Hook. f. New Zealand. 

L. leptoloba (Mattf.) D. Lloyd & C Webb. New Guinea. 
L. longipes Hook. f. Australia. 

L. maniototo (Petrie) D. Lloyd & C Webb. New Zealand. 
L. minor Hook. f. {Cotula haastii Kirk). New Zealand 
L. nana (D. Lloyd) D. Lloyd & C Webb. New Zealand. 
L. pectinata (Hook, f.) D. Lloyd & C Webb {Cotula monti- 

cola Simpson, C. villosa Simpson, C. willcoxii Cheese- 
man). New Zealand. 
L. plumosa Hook. f. New Zealand. 
L. potentillina (F. Muell.) Druce. New Zealand. 
L. pusilla Hook. f. {Cotula perpusilla Hook. f.). New 

Zealand. 
L. pyrethrifolia (Hook, f.) D. Lloyd & C Webb {Cotula 

linearifolia Cheeseman). New Zealand. 
*L. reptans (Benth.) D. Lloyd & C Webb. Austraha. 
L. rotundata (Cheeseman) D. Lloyd «fe C Webb. New 

Zealand. 
*L. sarawaketensis (P. Royen & D. Lloyd) D. Lloyd & C 

Webb. New Guinea. 
L. scariosa (Cass.) Franchet. Southern S. America, Austra- 
lia. 
L. serrulata (D. Lloyd) D. Lloyd & C Webb. New Zealand. 
L. squalida Hook. f. New Zealand. 
L. tenella (Cunn.) D. Lloyd & C Webb {Cotula membrana- 

cea D. Lloyd). New Zealand. 
L. traillii (Kirk) D. Lloyd & C Webb. New Zealand. 
L. wilhelminensis (P. Royen) D. Lloyd & C Webb. New 

Guinea. 

105. SOLIVA Ruiz Lopez & Pavon, Fl. peruv. prodr.: 
113 (1794). Type species: S. sessilis Ruiz Lopez & 
Pavon - Gymnostyles Juss. 

Annual herbs. Leaves alternate, pinnatisect. Capitula soli- 
tary, sessile in the leaf axils, disciform. Receptacle convex or 
conical, epaleate. Outer florets female, fertile, in several 
rows; tube and limb absent; style persistent and spinescent in 
fruit. Central florets functionally male (style-branches fused); 
corolla 3- or 4-lobed. Cypselas dorsiventrally flattened, with 2 
lateral vascular strands, laterally winged; wings sometimes 
projected to apical teeth, sometimes transversely rugose to 
sulcate. Pappus absent. 

Distribution. Mainly S. America but also in Australia and 
N. America, two species widespread as weeds {S. sessilis, S. 
stolonifera) . - 8 spp. 



160 



K. BREMER AND C. J. HUMPHRIES 



Soliva with Gymnostyles included is a well-defined mono- 
phyletic group characterized by the sessile capitula with 
female florets in several rows and functionally male central 
florets, and the persistent and more or less spinescent style. 
Gymnostyles, with the single species G. stolonifera cannot be 
retained (e.g. Tutin in Tutin et al., 1976). It differs from 
Soliva by its villous cypselas with transversely sulcate wings. 
Similar pubescent cypselas but with less extremely sulcate 
wings occur in some species of Soliva (5. anthemifolia) 
(Webb, 1986). The alternative would be to transfer 5. 
anthemifolia and related species to Gymnostyles but it seems 
quite unnecessary with two small, very similar and closely 
related genera rather than a single well-defined one. This 
solution is adopted also by Cabrera (1949) in his revision of 
the genus. 

5. anthemifolia R. Br. 
*S. macrocephala Cabrera 
*S. mutisii Kunth 
*5. neglecta Cabrera 

5. sessilis Ruiz Lopez «fe Pavon (S. pterosperma (Juss.) Less.) 
5. stolonifera (Brot.) R. Br. ex G. Don in Loudon {Gymno- 
styles stolonifera (Brot.) Tutin) 
*5. triniifolia Griseb. 
S. valdiviana Philippi 

106. SCHISTOSTEPHIUM Less, in Syn. gen. 
Compos.: 251 (1832). Type species: S. flabelliforme 
Less. - Peyrousea DC. 

Shrublets or half-shrubs. Leaves alternate, pinnatisect, lobed, 
dentate or occasionally entire. Capitula corymbose, occasion- 
ally solitary, disciform or discoid. Receptacle convex or 
conical, epaleate. Outer female florets tubular without limb, 
fertile, generally stalked. Central corolla 4-lobed. Cypselas 
dorsiventrally flattened, with 2 lateral (occasionally 3 or 4) 
vascular strands, sometimes with myxogenic cells. Pappus 
absent. 

Distribution. S. Africa from the E. Cape, Natal, and 
Transvaal to Mocambique and Zimbabwe. - 12 spp. 

Schistostephium is distinguished from the related genus Hip- 
pia by its 4-lobed corolla and perfect disc-florets. It was 
revised by Hutchinson (19166). The monotypic Peyrousea, 
revised by Bremer (1977, 1978), is very close to some species 
within Schistostephium, which appears to be paraphyletic 
with Peyrousea excluded. Hence we have merged the two 
genera. 

5. crataegifolium (DC.) Fenzl ex Harvey (5. villosum Hutch.) 

5. dactyliferum Hutch. 

5. flabelliforme Less. 

S. griseum (Harvey) Hutch. 

S. heptalobum (DC.) Oliver & Hiern (S. saxicola Hutch.) 

5. hippiifolium (DC.) Hutch. 

S. mollissimum Hutch. 

S. oxylobum S. Moore 

*S. rogersii Hutch. 

S. rotundifolium (DC.) Fenzl ex Harvey 

5. scandens Hutch. 

S. umbellatum (L. f.) Bremer & Humphries, comb. nov. 

Basionym: Cotula umbellata L. f., Suppl. pi.: 378 (1781) 

(Peyrousea umbellata (L. f.) Fourc). 



107. HIPPIA L., Mant. pi. 2: 158 (1771). Type 
species: H. frutescens (L.) L. 

Shrublets or half-shrubs. Leaves alternate, pinnatisect, lobed, 
dentate or occasionally entire. Capitula corymbose, occasion- 
ally solitary, disciform. Receptacle convex or conical, epale- 
ate. Outer female florets tubular without limb, fertile; tube 
much reduced. Central florets functionally male (style- 
branches fused); corolla 5-lobed, sometimes zygomorphic 
with 2 larger and 3 smaller lobes. Cypselas dorsiventrally 
flattened, with 2 lateral vascular strands, generally laterally 
winged. Pappus absent. 

Distribution. S. Africa in the Cape. - 8 spp. 

Hippia is traditionally distinguished by its functionally male 
central florets. It also differs by its 5-iobed corolla from the 
related Cotula and Schistostephium with 4-lobed corollas. In 
these two characters Hippia agrees with Eriocephalus but the 
two genera are otherwise very different. Hippia is generally 
treated together with Schistostephium, while Eriocephalus is 
considered isolated. For reasons of parsimony Eriocephalus is 
placed here as the sister group of Hippia. 
, Hippia was revised by Hutchinson (1918). H. montana and 
H. hutchinsonii were later described by Compton (1940) and 
Merxmiiller (Suessenguth and Merxmiiller, 1951), respec- 
tively. 

H. bolusae Hutch. 

H. frutescens (L.) L. 

H. hirsuta DC. 

*H. hutchinsonii Merxm. 

H. integrifolia Less. 

*H. montana Compton 

H. pilosa (P. J. Bergius) Druce 

*H. trilobata Hutch. 

108. ERIOCEPHALUS L., Sp. pi.: 926 (1753). Type 
species: E. africanus L. 

Shrubs, sometimes spinescent. Leaves alternate, sometimes 
opposite, often fascicled, variously lobed or entire, mostly 
ericoid. Capitula corymbose (pseudoumbellate or pseudora- 
cemose) or close together or occasionally solitary, radiate or 
disciform. Involucral bracts in 2 unequal series; one outer 
row of pubescent to glabrescent, scarious bracts and one 
inner row of generally densely villous, often connate bracts. 
Receptacle paleate; paleae generally villous. Ray or outer 
female florets fertile; Hmb white or mauve, short and wide, or 
absent. Disc florets functionally male (style-branches fused); 
corolla 5-lobed, yellow or mauve. Cypselas dorsiventrally 
flattened, with 2 lateral ribs with vascular strands. Pappus 
absent. Dehydrofalcarinone and dehydrofalcarinol present. 

Distribution. S. Africa mainly in the Cape but also in 
Namibia. - 26 spp. 

Eriocephalus is one of the most distinct and specialized 
genera in the tribe. The flattened fruits indicate a relationship 
with Cotula and its South African relatives Hippia and 
Schistostephium or possibly with Achillea and its relatives. It 
is here placed with the former group, since it agrees with 
Cotula in chemistry. Within the Cotula group Eriocephalus is 
most parsimoniously placed as sister group of Hippia, despite 
that Eriocephalus seems very different. 
The list of species is taken from Harvey (1865) with species 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 

described later added (e.g. Nordenstam, 1964). For species 
occurring in Namibia Merxmiiller (1967) should be consulted. 
The genus is presently under revision at Windhoek in 
Namibia. 

E. africanus L. 

E. aromaticus C. A. Smith 

E. aspalathoides DC. 

E. capitellatus DC. 

E. dinteri S. Moore 

E. ericoides (L. f.) Druce (£. glaber T\\\xnh .) 

E. eximius DC. 

*E. kingesii Merxm. & Eberle 

E. macroglossus B. Nord. 

E. microcephalus DC. 

*E. pauperrimus Merxm. & Eberle 

E. petrophiloides DC. 

E. pinnatus O. Hoffm. 

*E. pteronioides DC. 

E. pubescens DC. 

E. punctulatus DC. 

E. racemosus L. 

E. scariosissimus S. Moore 

E. scariosus DC. 

E. septulifer DC. 

E. sericeus Gaudich. ex DC. 

E. spinescens Burch. 

*E. tenuipes C. A. Smith 

E. tuberculosus DC. 

E. umbellulatus Cass. 

E. xerophilus Schltr 



EXCLUDED GENERA 

Potential future transfers, where known, are suggested in 
parenthesis. 

Abrotanella Cass. (Senecioneae - Blennospermatinae Less.) 

Baileya A. Gray (Heliantheae; near Psilostrophe DC.) 

Centipeda Lour (Astereae) 

Ceratogyne Turcz. (Astereae) 

Chondropyxis D. Cooke (Gnaphalieae) 

Dimorphocoma F. Muell. & Tate (Astereae) 

Elachanthus F. Muell. (Astereae) 

Formania W. W. Smith & Small (Astereae) 

Ischnea F. Muell. (Senecioneae - Blennospermatinae Less.) 

Isoetopsis Turcz. (Gnaphalieae) 

Lepidostephium Oliver (Gnaphalieae - Athrixiinae) 

Leucampyx A. Gray (Heliantheae; = Hymenopappus L' 

Herit.) 
Oedera L. (Gnaphalieae, near Relhania L' Herit. emend 

Anderb. & Bremer, and Ley sera L.) 
Plagiocheilus Am. ex DC. (Astereae; near Lagenophora 

Cass. & Solenogyne Cass.). 
Pseudocadiscus Lisowski (Senecioneae; = Stenops B. Nord.). 



Acknowledgements. This work would not have been possible 
without the support and assistance from botanical colleagues and 
fellow synantherologists. We would especially like to thank Dr Bertil 
Nordenstam and Mr John Cannon for providing working facilities in 
our respective institutions. We appreciate the assistance and 
co-operation of the directors of the following institutions for loaning 



161 

material or by giving us access to their collections (Abbreviations as 
in Holmgren et al., 1981): BM, BOL, BRY, E, F, K, GH, LE, M, 
MO, NBG, NY, PRE, S and US. Very special thanks to Mari 
Kallersjo for solutions to a great variety of taxonomic problems, 
especially of South African taxa. Many thanks also to Y. R. Ling who 
gave much assistance on the Artemisiinae tribe during his sabbatical 
year in the Department of Botany at the Natural History Museum 
during 1985-86 and to Helen B. Wilcox on the Anthemidineae and 
Chrysantheminae during her studies at Reading University. We are 
very grateful to Xi Shuh who provided us with helpful comments, 
literature, and specimens of Chinese taxa. Jens Klackenburg, at the 
University of Stockholm, gave much assistance in translating Russian 
botanical papers for which we are eternally grateful. We are grateful 
to Sylvia Gould for preparing the original index and for curating all of 
the collections at BM. Also, many minor errors would have remained 
undiscovered were it not for the diligent checking of species names 
and their synonyms by Sylvia Gould. Our thanks to Drs Norman 
Robson and Charlie Jarvis helping us on matters of nomenclature 
and botanical Latin. Thanks also to our reviewers Charles Jeffrey, 
Nicholas Hind and Fritz Ehrendorfer for their extremely helpful 
comments, although they should in no way be held responsible for 
the views we outline here. Finally, KB would like to thank the 
Swedish National Science Research Council for their generous finan- 
cial support. 



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INDEX 



Accepted names are in roman, synonyms in italics, new names and principal references in bold. 



Aaronsohnia Warb. & Eig 77, 143, 149, 150, 153, Achillea 

156, 157 Achillea 

Aaronsohnia factorovskyi Warb. & Eig 156, 157 Achillea 

Aaronsohnia pubescens (Desf.) Bremer & Achillea 

Humphries 157 Achillea 

Abrotanum Duhamel 120 Achillea 

Abrotanella Cass. 76, 161 Achillea 

Absinthium Miller 120 Achillea 

Achillea L. 77, 83, 106, 126, 127, 128, 129, 157, Achillea 

160 Achillea 
Achillea sect. Santolinoidea DC. 130 128 

Achillea abrotanoides (Vis.) Vis. 128 Achillea 

Achillea absinthoides Hal. 128 Achillea 

Achillea acuminata (Ledeb.) Schultz-Bip. 128 Achillea 

Achillea aegyptiaca L. 128 Achillea 

Achillea ageratifolia (Smith in Sibth. & Smith) Achillea 

Boiss. 128 128 



ageratum L. 128 Achillea 

aleppica DC. 128 Achillea 

alpina L. 128 Achillea 

ambrosiaca (Boiss. & Heldr.) Boiss. 128 Achillea 

arabica Kotschy 128 Achillea 

asiatica Serg. 128 Achillea 

asplenifolia Vent. 128 Achillea 

atrata L. 128 Achillea 

aucheri Boiss. 128 Achillea 

barbeyana Heldr. & Heimerl in Heimerl Achillea 

Achillea 

barrelieri (Ten.) Schultz-Bip. 106, 128 Achillea 

biebersteinii Afan. 128 Achillea 

biserrata M. Bieb. 128 Achillea 

boissieri (Hausskn.) Boiss. 128 Achillea 

brachyphylla Boiss. & Hausskn. in Boiss. Achillea 

Achillea 



bucharica Winkler 128 

caUichroa Boiss. 129 

camtschatica Rupr. ex Heimerl 129 

cappadocica Hausskn. & Bornm. 129 

cartilaginea Ledeb. ex Reichenb. 129 

chamaemelifolia Pourret 129 

chrysocoma Friv. 129 

clavennae L. 129 

clypeolata Sibth. & Smith 129 

coarctata Poiret in Lam. 129 

collina J. Becker ex Reichenb. 129 

compacta Willd. 129 

conferta DC. 129 

cretica L. 129 

crithmifolia Waldst. & Kit. 129 

cucullata (Hausskn.) Bornm. 129 

cuneatiloba Boiss. & Buhse 129 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



165 



Achillea decolorans Schrader 129 

Achillea depressa Janka 129 

Achillea distans Waldst. & Kit. ex Willd. 129 

Achillea erba-rotta All. 129 

Achillea falcata L. 129 

Achillea filipendulina Lam. 129 

Achillea fraasii Schultz-Bip. 129 

Achillea fragrantissima (Forssk.) Schultz-Bip. 129 

Achillea gerberi Willd. 129 

Achillea glaberrima Klokov 129 

Achillea goniocephala Boiss. & Bal. in Boiss. 129 

Achillea grandifolia Friv. 129 

Achillea griseo-virens Albov 129 

Achillea gypsicola Huber-Mor. 129 

Achillea haussknechtii Boiss. 129 

Achillea holosericea Sibth. & Smith 129 

Achillea impatiens L. 129 

Achillea inundata Kondr. in Wissjul. 129 

Achillea japonica Heimerl 129 

Achillea kellalensis Boiss. & Hausskn. in Boiss. 

129 
Achillea kermanica Gand. 129 
Achillea kotschyi Boiss. 129 
Achillea lanulosa Nutt. 129 
Achillea latiloba Ledeb. ex Nordm. 129 
Achillea ledebourii Heimerl 129 
Achillea leptophylla M. Bieb. 129 
Achillea ligustica All. 129 
Achillea hngulata Waldst. & Kit. 129 
Achillea lucana Pign. 129 

Achillea lycaonica Boiss. & Heldr. in Boiss. 129 
Achillea macrocephala Rupr. 129 
Achillea macrophylla L. 129 
Achillea magnifica Huber-Mor. 129 
Achillea maura Humbert 129 
Achillea membranacea (Labill.) DC. 129 
Achillea micrantha Willd. 128 
Achillea micranthoides Klokov 128 
Achillea millefolium L. 128, 129 
Achillea monocephala Boiss. & Bal. in Boiss. 129 
Achillea multifida (DC.) Boiss. 129 
Achillea nana L. 129 
Achillea neilrichii A. Kerner 129 
Achillea nobihs L. 129 
Achillea ochroleuca Ehrh. 129 
Achillea odorata L. 129 
Achillea oligocephala DC. 129 
Achillea oxylepis Boiss. & Hausskn. in Boiss. 129 
Achillea oxyloba (DC.) Schultz-Bip. 106 
Achillea oxyodonta Boiss. 129 
Achillea pachycephala Rech. f. 129 
Achillea pannonica Scheele 129 
Achillea phrygia Boiss. & Bal. in Boiss. 129 
Achillea pindicola Hausskn. 129 
Achillea pseudoaleppica Huber-Mor. 129 
Achillea ptarmica L. 129 
Achillea ptarmicifolia (Willd.) Rupr. ex Heimerl 

129 
Achillea ptarmicoides Maxim. 129 
Achillea pyrenaica Sibth. ex Godron in Gren. & 

Godron 129 
Achillea roseo-alba Ehrend. 129 
Achillea sachokiana Sosn. 129 
Achillea salicifoha Besser 129 
Achillea santolina L. 129 
Achillea santolinoides Lagasca 129 
Achillea schischkinii Sosn. 129 
Achillea schurii Schultz-Bip. 129 
Achillea sedelmeyeriana Sosn. 129 
Achillea septentrionalis (Serg.) Botsch. 129 
Achillea serbica Nyman 129 
Achillea setacea Waldst. & Kit. 129 
Achillea sibirica Ledeb. 129 
Achillea sieheana Stapf. 129 
Achillea sintenisii Huber-Mor. 129 
Achillea sipikorensis Hausskn. & Bornm. 129 
Achillea spinulifolia Fenzl ex Boiss. 129 
Achillea stricta (Koch) Schleicher ex Gremli 129 



Achillea sudetica Opiz 129 

Achillea talagonica Boiss. 129 

Achillea tanacetifolia All. 129 

Achillea taygetea Boiss. & Heldr. in Boiss. 129 

Achillea tenuifoha Lam. 129 

Achillea teretifolia Willd. 129 

Achillea thracica Velen. 129 

Achillea tomentosa L. 129 

Achillea umbellata Sibth. & Smith 129 

Achillea vermicularis Trin. 129 

Achillea virescens (Fenzl) Heimerl in A. Kerner 

129 
Achillea wilhelmsii Koch 129 
Achillea wilsoniana Heimerl ex Hand.-Mazz. 129 
Achilleinae Bremer & Humphries 74, 76, 77, 106, 

111, 126-128, 131, 137, 149 
Adenanthellum B. Nord. 82, 144, 145, 146 
Adenanthellum osmitoides (Harvey) B. Nord. 145, 

146 
Adenanthemum B. Nord. 146 
Adenoglossa B. Nord. 82, 147, 157 
Adenoglossa decurrens (Hutch.) B. Nord. 157 
Adenosolen DC. 152 
Adenosolen tenuif alius DC. 152 
AjaniaPolj.77, 81, 106, 110-112, 114, 115, 116 
Ajania achilleoides (Turcz.) Polj. ex Grubov 115 
Ajania adenantha (Diels) Ling & Shih 115 
Ajania aureoglobosa (W. Smith & Farrer) Muld. 

115 
Ajania brachyantha Shih 115 
Ajania breviloba (Franchet ex Hand.-Mazz.) Ling 

& Shih 115 
Ajania elegantula (W. Smith) Shih 115 
Ajania fastigiata (Winkler) Polj. 115 
Ajania fruticulosa (Ledeb.) Polj. 115, 116 
Ajania gracilis (Hook f. & Thomson) Polj. ex 

Tzvelev 115 
Ajania grubovii Muld. 115 
Ajania junnanica Polj. 115 
Ajania khartensis (Dunn) Shih 115 
Ajania kokanica (H. Kraschen.) Tzvelev 115 
Ajania latifolia Shih 115 
Ajania manshurica Polj. 116 
Ajania mutellina (Hand.-Mazz.) Muld. 115 
Ajania myriantha (Franchet) Ling ex Shih 115 
Ajania nana (H. Kraschen.) Muld. 115 
Ajania nematoloba (Hand.-Mazz.) Ling & Shih 

115 
Ajania nitida Shih 115 
Ajania nubigena (Wallich) Shih 115 
Ajania oresbia (W. Smith) Muld. 115 
Ajania pacifica (Nakai) Bremer & Humphries 115 
Ajania pallasiana (Fischer ex Besser) Polj. 115 
Ajania parviflora (Griin.) Ling 115 
Ajania potaninii (H. Kraschen.) Polj. 115 
Ajania przewalskii Polj. 115 
Ajania purpurea Shih 115 
Ajania quercifolia (W. Smith) Ling & Shih 116 
Ajania ramosa (Chang) Shih 116 
Ajania remotipinna (Hand.-Mazz.) Ling & Shih 

115 
Ajania roborowskii Muld. 115 
Ajania rupestris (Matsum. ex Koidz.) Muld. 115 
Ajania salicifolia (Mattf.) Polj. 116 
Ajania scharnhorstii (Regel & Schmalh.) Tzvelev 

115 
Ajania sericea Shih 115 
Ajania shiwogiku (Kitam.) Bremer & Humphries 

115 
Ajania tenuifolia (Jacquem.) Tzvelev 115 
Ajania tibetica (Hook. f. & Thomson) Tzvelev 115 
Ajania trilobata Polj. 115 
Ajania tripinnatisecta Ling & Shih 115 
Ajania variifolia (Chang) Tzvelev 116 
Ajaniopsis Shih 112, 116 
Ajaniopsis penicilliformis Shih 116 
AUardia Decne 81, 96, 97, 98, 99 
Allardia glabra Decne 98 



Allardia huegelii Schultz-Bip. 98 

Allardia lasiocarpa (G. X. H. C. Fu) Bremer & 

Humphries 98 
Allardia nivca Hook. f. & Thomson ex C. B. 

Clarke 98 
Allardia stoliczkae C. B. Clarke 98 
Allardia tomentosa Decne 98 
Allardia transalaica (Tzvelev) Bremer & 

Humphries 98 
Allardia tridactylites (Karehn & Kir.) Schultz-Bip. 

98 
Allardia vestita Hook. f. & Thomson ex C. B. 

Clarke 98 
Ambrosineae 75 

Ammanthus Boiss. & Heldr. ex Boiss. 132 
Anacyclus L. 77, 106, 126, 127, 129, 130 
Anacyclus clavatus (Desf.) Pers. 130 
Anacyclus cyrtolepidiodes Pomel 130 
Anacyclus homogamos (Maire) Humphries 130 
Anacyclus inconstans Pomel 130 
Anacyclus latealatus Huber-Mor. 130 
Anacyclus linearilobus Boiss. & Reuter 130 
Anacyclus maroccanus (Ball) Ball 130 
Anacyclus monanthos (L.) Thell. 130 
Anacyclus nigellifolius Boiss. 130 
Anacyclus officinarum Hayne 130 
Anacyclus pyrethrum (L.) Lagasca 130 
Anacyclus radiatus Lois. 130 
Anacyclus valentinus L. 129 
Anthemidaceae 75 
Anthemideae Cass. 73, 75-77, 80-83, 90, 91, 111, 

134, 144 
Anthemidinae Dumort. emend. Bremer & 

Humphries 74, 76, 77, 83, 131, 132, 137 
Anthemidees 75 
Anthemis L. 75, 77. 83, 108, 111, 131, 132-134, 

137, 139 
Anthemis L. sect. Maruta (Cass.) Boiss. 132 
Anthemis sect. Sclerorhachis Rech. f. 110 
Anthemis subgenus Ammanthus (Boiss. & Heldr.) 

R. Fernandes 75, 134 
Anthemis subgenus Anthemis 132 
Anthemis aaronsohnii Eig 132 
Anthemis abagensis Fed. 132 
Anthemis abrotanifolia (Willd.) Guss. 132 
Anthemis aciphylla Boiss. 132 
Anthemis adonidifolia Boiss. 132 
Anthemis aeolica Lojac. 132 
Anthemis aetnensis Schouw in Sprengel 132 
Anthemis alpestris (Hoffsgg & Link) R. Fernandes 

132 
Anthemis altissima L. 132 
Anthemis amblyolepis Eig 132 
Anthemis ammanthus Greuter 132 
Anthemis ammophila Boiss. & Heldr. in Boiss. 132 
Anthemis anahytae Woronow ex Sosn. 133 
Anthemis anatolica Boiss. 132 
Anthemis anthemiformis (Freyn & Sint.) Grierson 

132 
Anthemis antilibanotica Eig 132 
Anthemis antitaurica Grierson 132 
Anthemis arenicola Boiss. 132 
Anthemis argyrophylla (Hal. & Georgiev) Velen. 

132 
Anthemis armeniaca Freyn & Sint. 132 
Anthemis arvensis L. 132 
Anthemis atropatana Iranshar 132 
Anthemis auriculata Boiss. 132 
Anthemis austriaca Jacq. 132 
Anthemis austro-iranica Rech. f., Aellen & 

Esfand. 132 
Anthemis behboudiana Rech. f. & Esfand. 133 
Anthemis bornmuelleri Stoy. & Acht. 132 
Anthemis bourgaei Boiss. & Reuter 132 
Anthemis boveana Gay 132 
Anthemis brachmannii Boiss. & Heldr. in Boiss. 

132 
Anthemis brachycarpa Eig 132 



166 



K. BREMER AND C. J. HUMPHRIES 



Anthemis brachycentros Gay ex W. Koch 134 

Anthemis brachystephana Bornm. & Gauba 132 

Anthemis brevicuspis Bornm. 132 

Anthemis breviradiata Eig 132 

Anthemis bulgarica N.N. Thin 132 

Anthemis bushehrica Iranshahr 132 

Anthemis calcarea Sosn. 132 

Anthemis candidissima Willd. ex Sprengel 132 

Anthemis carpatica Waldst. & Kit. ex Willd. 132 

Anthemis caulescens Aitch. & Hemsley 110 

Anthemis chia L. 132 

Anthemis chrysantha Gay 132 

Anthemis coelopoda Boiss. 132 

Anthemis cornucopiae Boiss. 132 

Anthemis corymbulosa Boiss. & Hausskn. in 

Boiss. 133 
Anthemis cotula L. 132 
Anthemis cretacea Zefirov 133 
Anthemis cretica L. 133 
Anthemis cuneata Huber-Mor. & Reese 133 
Anthemis cypria Boiss. 133 
Anthemis cyrenaica Cosson 133 
Anthemis damascena Boiss. & Gaill. 133 
Anthemis davisii Yavin 133 
Anthemis debilifoha Eig 133 
Anthemis debilis Fed. 134 
Anthemis deserti Boiss. 133 
Anthemis deserticola H. Kraschen. & Popov 133 
Anthemis deserti-syriaci Eig 133 
Anthemis didymaea Mout. 133 
Anthemis dipsacea Bornm. 133 
Anthemis dubia Steven 133 
Anthemis dumetorum Sosn. 134 
Anthemis edumea Eig 133 
Anthemis eliezrae Eig 133 
Anthemis emasensis Eig 133 
Anthemis emiliae Sosn. 133 
Anthemis euxina Boiss. 134 
Anthemis feinbruniae Eig 132 
Anthemis filicaulis (Boiss. & Heldr.) Greuter 133 
Anthemis fimbriata Boiss. 133 
Anthemis flexicaulis Rech. f. 133 
Anthemis freitagii Iranshahr 133 
Anthemis fruticulosa M. Bieb. 133 
Anthemis fulvida Grierson 133 
Anthemis fumariifolia Boiss. 133 
Anthemis fumarioides Hochst. 133 
Anthemis fungosa Boiss. & Hausskn. 133 
Anthemis fuscata Brot. 130 
Anthemis galilaea Eig 132 
Anthemis gaudium-solis Velen. 133 
Anthemis gayana Boiss. 133 
Anthemis gerardiana Jordan 133 
Anthemis gilanica Boiss. 133 
Anthemis gilletti Iranshahr 133 
Anthemis glaberrima (Rech. f.) Greuter 133 
Anthemis glareosa Durieu & Barratte 133 
Anthemis gracilis Iranshahr 133 
Anthemis grossheimii Sosn. 133 
Anthemis halophila Boiss. & Bal. in Boiss. 133 
Anthemis hamrinensis Iranshahr 133 
Anthemis handel-mazzettii Eig 133 
Anthemis haussknechtii Boiss. & Renter in Boiss. 

133 
Anthemis hebronica Boiss. & Kotschy 133 
Anthemis hemistephana Boiss. 133 
Anthemis hermonis Eig 133 
Anthemis hinkovae N. N. Thin 133 
Anthemis hirtella Winkler 133 
Anthemis homalolepis Eig 133 
Anthemis hyalina DC. 133 
Anthemis hydruntina Groves 133 
Anthemis iberica M. Bieb. 133 
Anthemis indurata Del. 133 
Anthemis ismelia Lojac. 133 
Anthemis jailensis Zefirov 133 
Anthemis jordanovii Stoy. & Acht. 133 
Anthemis kandaharica Iranshahr 133 



Anthemis karabaghensis Mikheev 133 

Anthemis khorassanica Rech. f. 134 

Anthemis kitanovii N.N. Thin 133 

Anthemis kitenensis N.N. Thin 133 

Anthemis kotschyana Boiss. 133 

Anthemis krugeriana Pampan. 133 

Anthemis kurdica Iranshahr 133 

Anthemis kuzmanovii N. N. Thin 133 

Anthemis laconica R. Franzen 133 

Anthemis leptophylla Eig 133 

Anthemis leucanthemifolia Boiss. & Blanchet 133 

Anthemis leucolepis Eig 133 

Anthemis linczevskyi Fed. 133 

Anthemis hthuanica (DC.) Besser ex Trautv. 133 

Anthemis lorestanica Iranshahr 133 

Anthemis lyonnetioides (Boiss. & Kotschy) Boiss. 

133 
Anthemis macedonica Boiss. & Orph. in Boiss. 133 
Anthemis macrantha Heuffel 133 
Anthemis macroglossa Sommier & Levier 133 
Anthemis maris-mortui Eig 133 
Anthemis maris-nigri Fed. 133 
Anthemis maritima L. 90, 131, 132, 133 
Anthemis markhotensis Fed. 133 
Anthemis marschalliana Willd. 133 
Anthemis mauritiana Maire & Sennen 133 
Anthemis mazandaranica Iranshahr 133 
Anthemis melampodina Del. 133 
Anthemis melanacme Boiss. & Hausskn. in Boiss. 

133 
Anthemis melanolepis Boiss. 133 
Anthemis melanoloma Trautv. 133 
Anthemis meteorica Hausskn. 133 
Anthemis micrantha Boiss. & Hausskn. 133 
Anthemis microcephala (Schrenk) B. Fedtsch. 133 
Anthemis microlepis Eig 133 
Anthemis microsperma Boiss. & Kotschy 133 
Anthemis mirheydari Iranshahr 133 
Anthemis mixta L. 130 
Anthemis moghanica Iranshahr 133 
Anthemis monantha Willd. 133 
Anthemis monilicosta Pomel 133 
Anthemis montana L. 133 
Anthemis muricata (DC.) Guss. 133 
Anthemis nabataea Eig 133 
Anthemis nobilis L. 130 
Anthemis odontostephana Boiss. 133 
Anthemis orbelica Pancic 133 
Anthemis orientalis (L.) Degen 133 
Anthemis oxylepis (Boiss.) Boiss. 133 
Anthemis palestina Renter in Boiss. 133 
Anthemis panachaica Hal. 133 
Anthemis parnassica (Boiss. & Heldr.) R. 

Fernandes 133 
Anthemis parnesia Boiss. & Heldr. in Boiss. 133 
Anthemis parviceps Dobrocz. & Fed. 133 
Anthemis parvifolia Eig 133 
Anthemis patentissima Eig 133 
Anthemis pauciloba Boiss. 133 
Anthemis pectinata (Bory & Chaub.) Boiss. & 

Reuter 133 
Anthemis pedunculata Desf. 133 
Anthemis peregrina L. 134 
Anthemis persepolitana Boiss. 133 
Anthemis persica Boiss. 133 
Anthemis pestalozzae Boiss. 133 
Anthemis pindicola Heldr. ex Hal. 133 
Anthemis plebeia Boiss. & Noe 133 
Anthemis plutonia Meikle 133 
Anthemis praecox Link 130 
Anthemis pseudocotula Boiss. 133 
Anthemis ptarmiciformis K. Koch 133 
Anthemis punctata Vahl 134 
Anthemis pungens Yavin 134 
Anthemis rascheyana Boiss. 134 
Anthemis rayatensis Eig 132 
Anthemis regis-borisii Stoy. & Acht. 134 
Anthemis repanda L. 139 



Anthemis retusa Del. 134 

Anthemis rhodensis Boiss. 134 

Anthemis rhodocentra Iranshahr 134 

Anthemis rigescens Willd. 134 

Anthemis rigida (Sibth. & Smith) Boiss. & Heldr. 

134 
Anthemis rosea Smith in Sibth. & Smith 134 
Anthemis rumelica (Velen.) Stoy. & Acht. 134 
Anthemis ruthenica M. Bieb. 134 
Anthemis sabulifolia Pomel 134 
Anthemis saguramica Sosn. 134 
Anthemis samuelssonii Rech. f. 134 
Anthemis sancti-johannis Turrill 134 
Anthemis saportana Albov 134 
Anthemis scaettae Pampan. 134 
Anthemis scariosa Banks & Sol. in Russell 134 
Anthemis schischkiniana Fed. 134 
Anthemis schizostephana Boiss. & Hausskn. 134 
Anthemis scopulorum Rech. f. 134 
Anthemis scrobicularis Yavin 134 
Anthemis secundiramea Biv. 134 
Anthemis segetalis Ten. 134 
Anthemis semiensis Pichi-Serm. 134 
Anthemis sibthorpii Griseb. 134 
Anthemis sintenisii Freyn 134 
Anthemis sosnovskyana Fed. 134 
Anthemis spruneri Boiss. & Heldr. in Boiss. 134 
Anthemis sterilis Steven 134 
Anthemis stiparum Pomel 134 
Anthemis straussii Bornm. 133 
Anthemis stribrnyi Velen. 134 
Anthemis subtinctoria Dobrocz. 134 
Anthemis susiana Nab. 134 
Anthemis syriaca Bornm. 133 
Anthemis talyschensis Fed. 134 
Anthemis taubertii Durieu & Barratte 134 
Anthemis tempskyana Freyn & Sint. 133 
Anthemis tenuicarpa Eig 134 
Anthemis tenuiloba (DC.) R. Fernandes 134 
Anthemis tenuiflora Gilli 133 
Anthemis tigrensis Gay ex A. Richards 132, 134 
Anthemis tinctoria L. 132, 134 
Anthemis tomentella Greuter 134 
Anthemis tomentosa L. 134 
Anthemis tranzcheliana Fed. 134 
Anthemis tricolor Boiss. 134 
Anthemis tricornis Eig 134 
Anthemis tripolitana Boiss. & Blanchet in Boiss. 

134 
Anthemis triumfettii (L.) DC. in Lam. & DC. 134 
Anthemis trotzkiana Claus ex Bunge 134 
Anthemis tuberculata Boiss. 134 
Anthemis tubicina Boiss. & Hausskn. in Boiss. 133 
Anthemis virescens Velen. 134 
Anthemis wallii Huber-Mor. & Reese 134 
Anthemis werneri Stoy. & Acht. 134 
Anthemis wettsteiniana Hand.-Mazz. 134 
Anthemis wiedemanniana Fischer & C. Meyer 134 
Anthemis woronowii Sosn. 134 
Anthemis xylopoda O. Schwarz 134 
Anthemis yemenensis Podl. 134 
Anthemis zephyrovii Dobrocz. 134 
Anthemis zoharyana Eig 134 
Anthemis zyghia Woronow 134 
Arctanthemum (Tzvelev) Tzvelev 110, 111, 114, 

115 
Arctanthemum arcticum (L.) Tzvelev 114 
Arctanthemum hultenii (A. & D. Love) Tzvelev 

114 
Arctanthemum integrifolium (Richardson) Tzvelev 

114 
Arctanthemum kurilense (Tzvelev) Tzvelev 114 
Arctotideae Cass. 75 
Argyranthemum Webb ex Schultz-Bip. 83, 136, 

139 
Argyranthemum adauctum (Link) Humphries 136 
Argyranthemum broussonetii (Pers.) Humphries 

136 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



167 



Argyranthemum callichrysum (Svent.) Humphries 

136 
Argyranthemum coronopifohum (Willd.) 

Humphries 136 
Argyranthemum dissectum (Lowe) Lowe 136 
Argyranthemum x escarrei (Svent.) Humphries 136 
Argyranthemum filifohum (Schultz-Bip.) 

Humphries 136 
Argyranthemum foeniculaceum (Willd.) Webb ex 

Schultz-Bip. 136 
Argyranthemum frutescens (L.) Schultz-Bip. 136 
Argyranthemum gracile Schultz-Bip. 136 
Argyranthemum haematomma (Lowe) Lowe 136 
Argyranthemum haouarytheum Humphries & 

Bramwell 136 
Argyranthemum hierrense Humphries 136 
Argyranthemum jacobiifolium Kunkel 136 
Argyranthemum lemsii Humphries 136 
Argyranthemum lidii Humphries 136 
Argyranthemum maderense (D. Don) Humphries 

136 
Argyranthemum pinnatifidum (L. f.) Lowe 136 
Argyranthemum sundingii Borgen 136 
Argyranthemum sventenii Humphries & Aldridge 

136 
Argyranthemum tenerifae Humphries 136 
Argyranthemum thalassophilum (Svent.) 

Humphries 136 
Argyranthemum webbii Schultz-Bip. 136 
Argyranthemum winteri (Svent.) Humphries 136 
Artemisia L. 76, 77, 81, 83, 110-118, 120-125, 126, 

127, 147 
Artemisia sect. Abrotanum Besser 117, 120, 121 
Artemisia sect. Absinthium DC. 120, 121 
Artemisia sect. Artemisia 120 
Artemisia sect. Dracunculus Besser 81, 113, 116, 

120, 121, 125 
Artemisia sect. Euartemisia A. Gray 120 
Artemisia sect. Seriphidium Besser 117 
Artemisia subgenus Artemisia Less. 121 
Artemisia subgenus Dracunculus (Besser) Rydb. 

121 
Artemisia abaensis Y. R. Ling & S. Y. Zhao 121 
Artemisia abrotanum L. 121 
Artemisia absinthium L. 121 
Artemisia abyssinica Schultz-Bip. 121 
Artemisia adamsii Besser 121 
Artemisia afghanica Rech. f. 121 
Artemisia afra Jacq. 121 
Artemisia aksaiensis Y. R. Ling 121 
Artemisia alaskana Rydb. 121 
Artemisia albicerata H. Kraschen. 121 
Artemisia aleutica Hulten 121 
Artemisia altaiensis H. Kraschen. 121 
Artemisia amygdalina Decne 121 
Artemisia andersiana Podl. 121 
Artemisia anethifolia G. Weber in Stechm. 121 
Artemisia anethoides Mattf. 121 
Artemisia angustissima Nakai 121 
Artemisia annua L. 121 
Artemisia anomala S. Moore 121 
Artemisia aquatica Lour. 121 
Artemisia arborescens L. 121 
Artemisia arctica Less. 121 
Artemisia arctisibirica Korobkov 121 
Artemisia argilosa Beetle 118 
Artemisia argyi A. Leveille & Vaniot. 121 
Artemisia argyrophylla Ledeb. 121 
Artemisia armcniaca Lam. 121 
Artemisia aschurbajewii C. Winkler 121 
Artemisia atlantica Cosson & Durieu 121 
Artemisia atrata Lam. 121 
Artemisia atrovirens Hand.-Mazz. 121 
Artemisia aucheri Boiss. 121 
Artemisia aurata V. Komarov 121 
Artemisia australis Less. 121 
Artemisia austriaca Jacq. 121 
Artemisia austro-himalayensis (Y. R. Ling & H. S. 



Puri) Y. R. Ling & H. S. Puri 121 Artemisi 
Artemisia austro-yunnanensis Ling & Y. R. Ling Puri 

121 Artemisi 

Artemisia avarica Minat. 121 Artemisi 

Artemisia baimaensis Y. R. Ling & Z. C. Z. Y. Artemisi 

Zhuo 121 Artemisi 

Artemisia banihalensis Kaul & Bakshi 121 Artemisi 

Artemisia bargusinensis Sprcngel 121 Artemisi 

Artemisia bejdemaniae Leonova 121 Artemisi 

Artemisia bicolor Rech. f. & Wagenitz 118 Artemisi 

Artemisia biennis Willd. 121 Artemis 

Artemisia bigelowii A. Gray 118 Artemis 

Artemisia blepharolepis Bunge 121 Artemis 

Artemisia borealis Pallas 121, 122 Artemis 

Artemisia borealo-siamensis Y. R. Ling 121 Artemis 

Artemisia brachyloba Franchet 122 Artemis 

Artemisia brachyphylla Kitam. 122 Artemis 

Artemisia bre vis Pampan. 122 Artemis 

Artemisia burmanica Pampan. 122 Artemis^ 

Artemisia caerulescens var. cretacea Fiori 118 Artemis. 

Artemisia caespitosa Ledeb. 122 Artemisi 

Artemisia californica Less. 120, 122 Artemisi 

Artemisia calophylla Pampan. 122 Artemisi 

Artemisia campbellii Hook. f. & Thomson 122 Artemisi 

Artemisia campestris L. 121, 122 Artemis, 

Artemisia camphorata Villars 122 Artemisi 

Artemisia canadensis (Besser) Less. 122 Artemisi 

Artemisia cannabifolia A. Leveille 122 Artemisi 

Artemisia cantabrica (Lainz) Lainz 122 Artemisi 

Artemisia capillaris Thunb. 122 Artemisi 

Artemisia carruthii Wood 122 Artemisi 
Artemisia caruifolia Buch.-Ham. in Roxb. 122 122 

Artemisia cashimirica Kaul & Bakshi 122 Artemisi 

Artemisia caucasica Willd. 122 Artemisi 

Artemisia chamaemelifolia Villars 122 Artemis: 

Artemisia chiajeana Kunze 122 Artemis 

Artemisia chiarugii Pampan. 122 Artemis 

Artemisia chienshanica Ling & W. W. Wang 122 Artemis: 

Artemisia chingii Pampan. 122 Artemis 

Artemisia chitachensis Cosson ex Battand. & Artemis 

Trabut 122 Artemis 
Artemisia chitralensis Vo6\. \\^ 122 

Artemisia chrysolepis Kitagawa 122 Artemis 

Artemisia conaensis Ling & Y. R. Ling 122 Artemis 

Artemisia congesta Kitam. 122 Artemis 

Artemisia copa Philippi 122 Artemis 

Artemisia coracina W. W. Wang 122 Artemis 

Artemisia crithmifolia L. 122 Artemis 

Artemisia cuspidata H. Kraschen. 122 Artemis 

Artemisia daghestanica H. Kraschen. & Pors. 122 Artemis 

Artemisia dahurica (Turcz.) Polj. 122 Artemis 

Artemisia dalai-lamae H. Kraschen. 122 Artemis 

Artemisia demissa H. Kraschen. 122 Artemis 

Artemisia densifolia Filat. 122 Artemis 

Artemisia depauperata H. Kraschen. 122 Artemis 

Artemisia desertorum Sprengel 122 Artemis 

Artemisia dimoana Popov 122 Artemis 

Artemisia disjuncta H. Kraschen. 122 Artemisi 

Artemisia divaricata (Pampan.) Pampan. 122 Artemisi 

Artemisia diversa Diels 122 Artemis 

Artemisia dolichocephala Pampan. 122 Artemis 

Artemisia douglasiana Besser in Hook. f. 122 Artemis 

Artemisia dracunculiformis H. Kraschen. 122 Artemis 
Artemisia dracunculus L. 122 123 

Artemisia dubia Wallich ex Besser 122 Artemisi 

Artemisia dudinensis V. P. Amel'chenko 122 Artemisi 

Artemisia duthreuil-de-rhinsi H. Kraschen. 122 Artemisi 

Artemisia edgeworthii Balakr. 122 Artemisi 

Artemisia eldarica Rzazade 122 Artemisi 

Artemisia elegantissima Pampan. 122 Artemisi 

Artemisia emeiensis (Chang) Y. R. Ling 122 Artemisi 

Artemisia eriantha Ten. 122 Artemisi 

Artemisia eriocephala Pampan. 122 Artemis, 

Artemisia eriopoda Bunge 122 Artemisi 

Artemisia eriangshanensis Ling & Y. R. Ling 122 Artemisi 

Artemisia faurieri Nakai 122 Artemisi 

Artemisia feddei A. Leveille & Vaniot. 123 Artemisi 

Artemisia filifolia Torrey 122 Artemisi 



a filiformilobulata Y. R. Ling & H. S. 
122 

a flaccida Hand.-Mazz. 122 

a flahaultii Emb. & Maire 122 

a flava Jurtzev 122 

a flavifolia Gilli 122 

a forrestii W. Smith 122 

a franserioides Greene 122 

a freyniana (Pampan.) H. Kraschen. 122 
a freitagii Pod\. 118 

a frigida Willd. 122 

a frigidioides H. C. Fu & Z. Y. Zhu 122 

a fukudo Makino 122 

a fulgens Pampan. 122 

a furcata M. Bieb. 122 

a gabriellae Braun-Blanquet 122 

a gangsuensis Ling & Y. R. Ling 122 

a genipi G. Weber in Stechm. 122 
'a ghazniensis Podl. 118 
'a ghoratensis Podl. 118 

a gilvescens Miq. 122 

a giraldii Pampan. 122 

a glabella Karelin & Kir. 122 

a glacialis L. 122 
la glauca Pallas ex Willd. 122 

a globosa H. Kraschen. 122 

a globosoides Ling & Y. R. Ling 122 

a globularia Cham, ex Besser 122 

a glomerata Ledeb. 122 

a gmelinii G. Weber in Stechm. 122 

a gongshanensis Y. R. Ling & Humphries 

a gorgonum Webb in Hook. 122 

a granatensis Boiss. 122 

a graveolens Minat. 122 

a gyangzeensis Ling & Y. R. Ling 122 

a gyitangensis Ling & Y. R. Ling 122 

a haichowensis Chang 122 

a hallaisanensis Nakai 122 

a halodendron Turcz. ex Besser 122 

a hancei (Pampan.) Ling & Y. R. Ling 

a haussknechtii Boiss. 122 

a hedinii Ostenf. & Pauls in Hedin 122 

a henriettae H. Kraschen. 122 

a hillebrandii Skottsb. 122 

a hippolytii Butkov 122 

a hispanica Lam. 122 

a hololeuca M. Bieb. ex Besser 122 

a hulteniana Vorosch. 122 

a hultenii Maksimova 122 

a idilongensis Y. R. Ling 122 

a ifranensis J. Didier 122 

a igniaria Maxim. 122 

a implicata Leonova 122 

a imponens Pampan. 122 

a incana (L.) Druce 122 

a incisa Pampan. 122 

a indica Willd. 123 

a insipida Villars 123 

a insulana H. Kraschen. 123 

a integrifolia L. 123 

a intramongolica H. C. Fu & Z. Y. Zhu 

la jacutica Drob. 123 
la japonica Thunb. 123 
la javanica Pampan. 123 

jaxatica Polj. 123 

jilongensis Y. R. Ling & Humphries 123 

judaica L. 123 

kabylica Chabert. 123 

kanashiroi Kitam. 123 
ia kandaharensis Podl. 119 

kangmasensis Ling & Y. R. Ling 123 

karavajevii Leonova 123 

kauaiensis (Skottsb.) Skottsb. 123 

kawakamii Hayata 123 

keiskeana Miq. 123 



168 



K. BREMER AND C. J. HUMPHRIES 



Artemisia kelleri H. Kraschen. 123 
Artemisia kermanensis Podl. 119 
Artemisia khorassanica Podl. 119 
Artemisia kitadakensis Hara & Kitam. 123 
Artemisia klementzae H. Kraschen. ex Leonova 

123 
Artemisia klotzschiana Besser 123 
Artemisia koidzumii Nakai 123 
Artemisia komarovii Polj. 123 
Artemisia kulbadica Boiss. & Buhse 123 
Artemisia kumykorum Minat. 123 
Artemisia kuschakewiczii Winkler 123 
Artemisia laciniatiformis V. Komarov 123 
Artemisia lactiflora Wallich ex DC. 123 
Artemisia lagocephala (Fischer ex Besser) DC. 123 
Artemisia lagopus Fischer ex Besser 123 
Artemisia lamprocaulos Rech. f. 123 
Artemisia lancea Vaniot 123 
Artemisia latifolia Ledeb. 123 
Artemisia lavandulifolia DC. 123 
Artemisia lavei Kostel. 123 
Artemisia ledebouriana Besser 123 
Artemisia leontopodioides Fischer ex Besser 123 
Artemisia leptophylla D. Don 123 
Artemisia leucophylla (Turcz. ex Besser) C. B. 

Clarke 123 
Artemisia limosa Koidz. 123 
Artemisia limprichtii (Pampan.) Ling & Y. R. Ling 

123 
Artemisia hpskyi Polj . 123 
Artemisia littoricola Kitam. 123 
Artemisia longifolia Nutt. 123 
Artemisia ludoviciana Nutt. 123 
Artemisia macilenta (Maxim.) H. Kraschen. 123 
Artemisia maciravae Hutch. & Dalziel 123 
Artemisia macrantha Ledeb. 123 
Artemisia macrocephala Jacq. 123 
Artemisia macrorhiza Turcz. 123 
Artemisia magellanica Schultz-Bip. 123 
Artemisia mairei A. Leveille 123 
Artemisia manshurica (V. Komarov) V. Komarov 

123 
Artemisia maritima L. ssp. kasakorum H. 

Kraschen. 119 
Artemisia maroccana Cosson 123 
Artemisia marschalliana Sprengel 123 
Artemisia martjanovii H. Kraschen. ex Polj. 123 
Artemisia mattfeldii Pampan. 123 
Artemisia mauiensis (A. Gray) Skottsb. 123 
Artemisia maximovicziana (F. Schum.) H. 

Kraschen. ex Polj. 123 
Artemisia medioxima H. Kraschen. ex Polj. 123 
Artemisia melanolepis Boiss. & Kotschy 123 
Artemisia mendozana DC 119 
Artemisia mesatlantica Maire 123 
Artemisia michauxiana Besser 123 
Artemisia minor Jacq. in Besser 123 
Artemisia molinieri Quezel, Barbero & R. Loisel 

123 
Artemisia moUuccana Roxb. 123 
Artemisia momiyamae Kitam. 123 
Artemisia mongolica (Fischer ex Besser) Nakai 123 
Artemisia monophylla Kitam. 123 
Artemisia monosperma Del. 123 
Artemisia montana Pampan. 123 
Artemisia montevidensis Sprengel 123 
Artemisia moorcroftiana WaUich ex DC. 123 
Artemisia morrisonensis Hayata 123 
Artemisia multisecta Leonova 123 
Artemisia mutellina Villars 123 
Artemisia myriantha Wallich ex DC. 123 
Artemisia nakaii Pampan. 123 
Artemisia nanshanica H. Kraschen. 123 
Artemisia neglecta Leonova 123 
Artemisia negrei Ouyahya 123 
Artemisia nesiotica Raven 123 
Artemisia niitakayamensis Hayata 123 
Artemisia nilagirica (C. B. Clarke) Pampan. 123 



Artemisia nitida Bertol. 123 

Artemisia nivalis Braun-Blanquet 123 

Artemisia nortonii Pampan. 123 

Artemisia norvegica Fries 123 

Artemisia nujianensis (Ling & Y. R. Ling) Y. R. 

Ling 123 
Artemisia nuristanica Kitam. 122 
Artemisia obscura Pampan. 123 
Artemisia obtusiloba Ledeb. 123 
Artemisia occidentali-sichuanensis Y. R. Ling & S. 

Y. Zhao 123 
Artemisia occidentah-sinensis Y. R. Ling 123 
Artemisia occidentaH-yunnanensis Ling & Y. R. 

Ling 123 
Artemisia oelandica (Besser) V. Komarov 123 
Artemisia olchonensis Leonova 123 
Artemisia olgensis (Vorobiev) Vorosch. 123 
Artemisia oligocarpa Hayata 123 
Artemisia opulenta Pampan. 123 
Artemisia oranensis Filat. 123 
Artemisia ordosica H. Kraschen. 123 
Artemisia orientahs (Pampan.) Ling & Y. R. Ling 

123 
Artemisia orientali-hengduangensis Ling & Y. R. 

Ling 123 
Artemisia orientali-xizangensis Y. R. Ling & 

Humphries 123 
Artemisia orientali-yunnanensis Y. R. Ling 123 
Artemisia orthobotrys Kitagawa 123 
Artemisia oxycephala Kitagawa 123 
Artemisia packardiae Grimes & Ertter 123 
Artemisia pallens Wallich ex Besser 123 
Artemisia palmeri A. Gray 119 
Artemisia palustris L. 123 
Artemisia pancicii (Janka) Ronniger 123 
Artemisia pannosa H. Kraschen. 123 
Artemisia papposa Blake & Cronq. 123 
Artemisia parryi A. Gray 124 
Artemisia parviflora Buch.-Ham. ex Roxb. 124 
Artemisia pattersonii A. Gray 124 
Artemisia pedatifida Nutt. 124 
Artemisia pedunculosa Miq. 124 
Artemisia pengchuoensis Y. R. Ling & S. Y. Zhao 

124 
Artemisia persica Boiss. 124 
Artemisia pewzowii Winkler 124 
Artemisia phaeolepis H. Kraschen. 124 
Artemisia phyllobotrys (Hand.-Mazz.) Ling & Y. 

R. Ling 124 
Artemisia polybotryoidea Y. R. Ling 124 
Artemisia pontica L. 124 
Artemisia ported Cronq. 124 
Artemisia praticola Klokov 124 
Artemisia prattii (Pampan.) Ling & Y. R. Ling 124 
Artemisia princeps Pampan. 124 
Artemisia przewalskii H. Kraschen. 124 
Artemisia pseudopontica Schur. 124 
Artemisia pubescens Ledeb. 124 
Artemisia punctigera H. Kraschen. 124 
Artemisia pycnorhiza Ledeb. 124 
Artemisia quettensis Podl. 119 
Artemisia quinlingensis Ling & Y. R. Ling 124 
Artemisia quinqueloba Trautv. 124 
Artemisia ramosa C. Smith 124 
Artemisia rehan Chiov. 124 
Artemisia remotiloba H. Kraschen. ex Polj. 124 
Artemisia reptans C. Smith ex Link 124 
Artemisia robusta (Pampan.) Ling & Y. R. Ling 

124 
Artemisia rosthornii Pampan. 124 
Artemisia roxburghiana Besser 124 
Artemisia rubripes Nakai 124 
Artemisia rupestris L. 124 
Artemisia rutifolia Stephen ex Sprengel 124 
Artemisia sacrorum Ledeb. 124 
Artemisia saitoana Kitam. 124 
Artemisia salsoloides Willd. 124 
Artemisia samoiedorum Pampan. 124 



Artemisia santohnifolia Turcz. ex H. Kraschen. 

124 
Artemisia saposhnikovii H. Kraschen. ex Polj. 124 
Artemisia schimperi Schultz-Bip. ex Schweinf. 124 
Artemisia schischkinii H. Kraschen. 124 
Artemisia schmidtiana Maxim. 124 
Artemisia scoparia Waldst. & Kit. 124 
Artemisia scopulorum A. Gray 124 
Artemisia selengensis Turcz. ex Besser 124 
Artemisia senjavinensis Besser 124 
Artemisia sericea G. Weber in Stechm. 124 
Artemisia serrata Nutt. 124 
Artemisia serreana Pampan. 124 
Artemisia shangnanensis Ling & Y. R. Ling 124 
Artemisia shennongjaensis Ling & Y. R. Ling 124 
Artemisia sichuanensis Ling & Y. R. Ling 124 
Artemisia sieversiana Ehrhart in Willd. 124 
Artemisia simulans Pampan. 124 
Artemisia sinanensis Yabe 124 
Artemisia sinensis (Pampan.) Ling & Y. R. Ling 

124 
Artemisia smithii Mattf. 124 
Artemisia somai Hayata 124 
Artemisia songarica Schrenk. 124 
Artemisia speciosa (Pampan.) Ling & Y. R. Ling 

124 
Artemisia sphaerocephala H. Kraschen. 124 
Artemisia spiciformis var. longiloba Osterh. 119 
Artemisia spinescens D. Eaton 125 
Artemisia splendens Willd. 124 
Artemisia stelleriana Besser 124 
Artemisia stenophylla Kitam. 124 
Artemisia stipularis Urb. & Ekman 124 
Artemisia stolonifera (Maxim.) V. Komarov 124 
Artemisia stracheyi Hook f. & Thomson ex C. B. 

Clarke 124 
Artemisia stricta Edgew. 124 
Artemisia subchrysolepis F\\ai. 119 
Artemisia subulata Nakai 124 
Artemisia subviscosa Turcz. 124 
Artemisia succulenta Ledeb. 124 
Artemisia succulentoides Ling & Y. R. Ling 124 
Artemisia suksdorfii Piper 124 
Artemisia superba Pampan. 124 
Artemisia swatensis Podl. 124 
Artemisia sylvatica Maxim. 124 
Artemisia tafelii Mattf. 124 
Artemisia taibaishanensis Y. R. Ling & Humphries 

124 
Artemisia tainingensis Hand.-Mazz. 124 
Artemisia tanacetifolia L. 124 
Artemisia tangutica Pampan. 124 
Artemisia tecti-mundii Podl. 119 
Artemisia tenuifolia Y. R. Ling & H. S. Puri 124 
Artemisia thellungiana Pampan. 124 
Artemisia tilesii Ledeb. 124 
Artemisia tomentella Trautv. 124 
Artemisia tournefortiana Reichenb. 124 
Artemisia transbaicalensis Leonova 124 
Artemisia trautvetteriana Besser 124 
Artemisia tridactyla Hand.-Mazz. 124 
Artemisia triniana Besser 124 
Artemisia tschernieviana Besser 124 
Artemisia tsugitakaensis (Kitam.) Ling & Y. R. 

Ling 124 
Artemisia tsuneoi Tatewaki & Kitam. 124 
Artemisia tukuchaensis Kitam. 124 
Artemisia tyitangensis Ling & Y. R. Ling 124 
Artemisia unalaskensis Rydb. 124 
Artemisia ussuriensis Polj. 124 
Artemisia velutina Pampan. 124 
Artemisia verbenacea (V. Komarov) Kitagawa 124 
Artemisia verlotorum Lamotte 124 
Artemisia vestita Wallich ex Besser 124 
Artemisia vexans Pampan. 124 
Artemisia viridisquama Kitam. 124 
Artemisia viridissima (V. Komarov) Pampan. 125 
Artemisia viscida (Mattf.) Pampan. 125 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



Artemisia viscidissima Ling & Y. R. Ling 125 

Artemisia vulgaris L. 110, 120, 125 

Artemisia waltonii J. R. Drumm. ex Pampan. 125 

Artemisia wellbyi Hemsley & Pears 125 

Artemisia wudanica Liou & W. Wang 125 

Artemisia xanthochloa H. Kraschen. 125 

Artemisia xcrophytica H. Kraschen. 125 

Artemisia xigazcensis Ling & Y. R. Ling 125 

Artemisia yadongensis Ling & Y. R. Ling 125 

Artemisia yongii Y. R. Ling 125 

Artemisia younghusbandii J. R. Drumm. 125 

Artemisia yunnanensis Jeffrey ex Dicls 125 

Artemisia zayuensis Ling & Y. R. Ling 125 

Artemisia zhondianensis Y. R. Ling 125 

Artemisiastrum Rydb. 117, 118 

Artemisiastrum palmeri {A. Gray) Rydb. 118, 119 

Artemisiella Ghafoor 120 

Artemisiella stracheyi (Hook. f. & Thompson ex C. 

B. Clarke) Ghafoor 124 
Artemisiinae Less, emend. Bremer & Humphries 
73, 74, 76, 77, 80-84, 110-113, 116, 117, 127, 132, 

137, 138, 139, 147 
Asaemia (Harvey) Harvey ex Benth. in Benth. & 

Hook. f. 92, 93, 95, 96 
Asaemia axillaris (Thunb.) Harvey ex Hoffmann 

96 
Asaemia minuta (L. f.) Bremer 96 
Asaemia minuta ssp. inermis (E. PhiUips) Bremer 

96 
Asaemia minuta (L. f.) Bremer ssp. minuta 96 
Asteraceae 73-75, 77, 81, 83 
Astereae 74, 90, 161 

Asteroideae 75 

Athanasia L. 77, 91-93, 94, 95, 96, 107, 108 

Athanasia acerosa (DC.) D. Dietr. 94, 95 

Athanasia adenantha (Harvey) Kallersjo 95 

Athanasia alba Kallersjo 95 

Athanasia bremeri Kallersjo 95 

Athanasia brownii Hochr. 108 

Athanasia calophylla Kallersjo 95 

Athanasia calva Hutch. 108 

Athanasia capitata (L.) L. 95 

Athanasia cochlearifolia Kallersjo 95 

Athanasia coronopifolia Harvey 108 

Athanasia crenata (L.) L. 95 

Athanasia crithmifoha (L.) L. 95, 95 

Athanasia cuneifolia Lam. 95 

Athanasia dentata (L.) L. 95 

Athanasia dregeana (DC.) Harvey 108 

Athanasia elsiae Kallersjo 96 

Athanasia filiformis L. f. 96 

Athanasia flexuosa Thunb. 96 

Athanasia grandiceps Hilliard & Burtt 95, 96 

Athanasia hirsuta Thunb. 96 

Athanasia humilis Kallersjo 95, 96 

Athanasia imbricata Harvey 96 

Athanasia inopinata (Hutch.) Kallersjo 96 

Athanasia juncea (DC.) D. Dietr. 96 

Athanasia leptocephala Kallersjo 96 

Athanasia leucoclada (DC.) Harvey 108 

Athanasia linifolia L. f. 96 

Athanasia microcephala (DC.) D. Dietr. 96 

Athanasia microphylla DC. 96 

Athanasia minuta (L. f.) Kallersjo 95, 96 

Athanasia montana J. M. Wood & M. Evans 108 

Athanasia oocephala (DC.) Kallersjo 96 

Athanasia pachycephala DC. 96 

Athanasia pectinata L. f. 96 

Athanasia pinnata L. f. 96 

Athanasia pinnatifida (Oliver) Hilliard 94 

Athanasia pubescens (L.) L. 96 

Athanasia punctata (DC.) Harvey 108 

Athanasia quinquedentata Thunb. 96 

Athanasia rugulosa E. Meyer ex DC. 96 

Athanasia scabra Thunb. 96 

Athanasia schistostephioides Hiern 108 

Athanasia schizolepis Harvey 95 

Athanasia scrtulifera DC. 96 



Athanasia spathulata (DC.) D. Dietr, 96 
Athanasia thodei Bolus 108 
Athanasia tomentosa Thunb. 96 
Athanasia tridens Oliver 108 
Athanasia trifurcata (L.) L. 96 
Athanasia vcstita (Thunb.) Druce 96 
Athanasia villosa Hilliard 94 
Athanasia virgata Jacq. 96 
Athanasia viridis Kallersjo 96 
Athanasia woodii (Thell.) Hilliard 94 
Athrixiinae 161 

Bailey a A. Gray 161 

Balsamita Miller 100, 101 

Balsamita major Desf. 102 

Blennospermatinae Less. 161 

Brachanthemum DC. 110, 113, 114 

Brachanthemum sect. Brachanthemum 113 

Brachanthemum sect. Dcndranthemopsis Tzvelev 

113 
Brachanthemum baranovii (H. Kraschen. & Polj.) 

H. Kraschen. 113 
Brachanthemum fruticulosum (Ledeb.) DC. 113 
Brachanthemum gobicum H. Kraschen. 113 
Brachanthemum kasakhorum H. Kraschen. 113 
Brachanthemum kirghisorum H. Kraschen. 113 
Brachanthemum krylovii Serg. 113 
Brachanthemum mongolicum H. Kraschen. 113 
Brachanthemum mongolorum Grubov 113 
Brachanthemum nanshanicum H. Kraschen. 113 
Brachanthemum pulvinatum (Hand.-Mazz.) Shih 

113 
Brachanthemum titovii H. Kraschen. 113 
Brachymeris DC. 94 

Brachymeris athanasioides (S. Moore) Hutch. 94 
Brachymeris bolusii Hutch. 94 
Brachymeris erubesceits Hutch. 94 
Brachymeris montana Hutch. 94 
Brachymeris peglerae Hutch. 94 
Brachymeris scoparia DC. 94 
Brocchia Vis. 157, 158 
Brocchia cinerea (del.) Vis. 158 

Calenduleae 74 

Cancrinia Karelin & Kir. 83, 96, 97, 98, 99, 101, 

104, 106, 154 
Cancrinia sect. Matricarioides Tzvelev 99 
Cancrinia sect. Polychrysum Tzvelev 109 
Cancrinia sect. Tanacetopsis Tzvelev 104 
Cancrinia botschantzevii (Kovalevsk.) Tzvelev 104 
Cancrinia discoidea (Ledeb.) Polj. ex Tzvelev 99, 

154 
Cancrinia chrysocephala Karelin & Kir. 96, 99 
Cancrinia ferganensis (Kovalevsk.) Tzvelev 104 
Cancrinia golovskovii (Polj.) Tzvelev 104 
Cancrinia karatavica Tzvelev 104 
Cancrinia krasnoborovii V. Khan 99 
Cancrinia maximoviczii Winkler 102 
Cancrinia mucronata (Regel & Schmalh.) 

Kovalevsk. 104 
Cancrinia nevskii Tzvelev 104 
Cancrinia pamiralaica (Kovalevsk.) Kovalevsk. 99 
Cancrinia pjataeviae (Kovalevsk.) Tzvelev 104 
Cancrinia santoana (H. Kraschen. Popov & Vved.) 

Tzvelev 104 
Cancrinia setacea (Regel & Schmalh.) Tzvelev 104 
Cancrinia submarginata (Kovalevsk.) Tzvelev 104 
Cancrinia subsimilis (Rech. f.) Tzvelev 104 
Cancrinia tianshanica (H. Kraschen.) Tzvelev 99 
Cancrinia urgutensis (Popov) Tzvelev 105 
Cancriniella Tzvelev 96, 97, 98. 99 
Cancriniella krascheninnikovii (Rubtzov) Tzvelev 

99 
Cancriniinae Bremer & Humphries 74, 76, 77, 81, 

83, 91,%, 97, 101, 138 
Cenia Comm. ex Juss. 157, 158 
Cenia albovillosa S. Moore 158 
Cenia duckittiae L. Bolus 158 



169 

Cenia expansa Compton 158 

Cenia microglossa DC. 158 

Cenia pectinata DC. 158 

Cenia sericea (L.f.) DC. 159 

Cenia turbinata (L.) Pers. 159 

Centipeda Lour 76, 161 

Ceratogy ne Tmcz. 76, 161 

Chamaemelum Miller 77, 126, 127, 130, 131 

Chamaemelum callosum Boiss. & Heldr. 156 

Chamaemelum daghestanicum Rupr. ex Boiss. 102, 

156 
Chamaemelum eriolepis (Cosson ex Maire) Benedi 

130 
Chamaemelum flahauiti (Emb.) Benedi 130 
Chamaemelum fuscatum (Brot.) Vase. 130 
Chamaemelum grandiflorum Boiss. & Hausskn. 

156 
Chamaemelum heterolepis Freyn & Sint. 156 
Chamaemelum kotschyi Boiss. 156 
Chamaemelum mixtum (L.) All. 130 
Chamaemelum nobile (L.) All. 130 
Chamaemelum repens Freyn & Sint. 
Chamaemelum scariosum (Ball) Benedi 130 
Chamartemisia Rydb. 117 

Chamartemisia compacta (H. M. Hall) Rydb. 117 
Chamomilla Gray 153, 154 
Chamomilla aurea (Loefl.) Gay ex Cosson & 

Kralik 
Chamomilla lasiocarpa (Boiss.) Rauschert 154 
Chamomilla macrotis (Rech. f.) Rauschert 154 
Chamomilla occidentalis (Greene) Rydb. 154 
Chamomilla pubescens (Desf.) Alavi 157 
Chamomilla suaveolens (Pursh) Rydb. 154 
Chamomilla tzvelevii (Pobed.) Rauschert 156 
Chamomilla vulgaris Gray 154 
Chlamydophora Ehrenb. ex Less. 139, 142, 143 
Chlamydophora pubescens (Desf.) Cosson & 

Durieu 143, 157 
Chlamydophora tridentata (del.) Ehrenb. ex Less. 

142, 143 
Chrysanthemeae Less 134 
Chondropyxis D. Cooke 161 
Chrysantheminae Less, emend. Bremer & 

Humphries 73-77, 81, 83, 84, 128, 131 
134, 135 
Chrysanthemum L. 76, 77, 83, 84, 110, 113, 134, 

142, 151 

Chrysanthemum afghanicum Gilli 104 
Chrysanthemum atlanticum Ball 142 
Chrysanthemum carinatum Schousboe 135 
Chrysanthemum catananche Ball 142 
Chrysanthemum chalchingolicum Grubov 114 
Chrysanthemum clausonis (Pomel) Battand. 143 
Chrysanthemum coronarium L. 84, 134, 135 
Chrysanthemum crassicollum Rech. f. 106 
Chrysanthemum cuneifolium Kitam. 114 
Chrysanthemum deserticola (Murb.) F. Buxbaum 

143, 153 

Chrysanthemum djilgense Franchet 98 
Chrysanthemum dolichophyllum Kitam. 106 
Chrysanthemum fontanesii (Boiss. & Reuter) 

Ouezel & Santa 141 
Chrysanthemum fuscatum Desf. 153 
Chrysanthemum gayanum Ball 142 
Chrysanthemum gayanum var. depressum Ball 142 
Chrysanthemum kelleri Krylov & Plotn. 102 
Chrysanthemum macrocarpum Cosson & Kralik ex 

Battand. in Battand. & Trab. 154 
Chrysanthemum maresii var. hosmariense Ball 142 
Chrysanthemum maroccanum Battand. 142 
Chrysanthemum marschallii Aschers. 101 
Chrysanthemum multicaule Desf. 143 
Chrysanthemum myconis L. 143 
Chrysanthemum nipponicum Franchet ex Maxim. 

139 
Chrysanthemum nivellei Braun-Blanquet & Maire 

140 
Chrysanthemum pacificum Nakai 115 



170 



K. BREMER AND C. J. HUMPHRIES 



Chrysanthemum paludosum Poiret 142 
Chrysanthemum porphyrostephanum Rech. f. 103 
Chrysanthemum richterioides Winkler 103 
Chrysanthemum segetum L. 84, 135 
Chrysanthemum shiwogiku Kitam. 115 
Chrysanthemum tatsienense Bureau & Franchet 103 
Chrysanthemum trifurcatum Desf. 143, 153 
Chrysanthemum viscidehirtum (Schott) Thell. 135 
Chrysanthoglossum Wilcox, Bremer & Humphries 

77, 138, 139, 143 
Chrysanthoglossum deserticola (Murb.) Wilcox, 

Bremer & Humphries 143 
Chrysanthoglossum trifurcatum (Desf.) Wilcox, 

Bremer & Humphries 143 
Cladanthus Cass. 126, 127, 130, 131 
Cladanthus arabicus (L.) Cass. 131 
Coleostephus Cass. 77, 136, 137, 139, 143. 144 
Coleostephus clausonis Pomel 143 
Coleostephus multicaulis (Desf.) Durieu 143 
Coleostephus myconis (L.) Reichenb. f. 143 
Coleostephus paludosus (Durieu) Alavi 143 
Cotula L. 75-77, 81, 83, 101, 127, 147-151, 155, 

157-159, 160 
Cotula sect. Cotula 158, 159 
Cotula sect. Leptinella (Cass.) Hook. f. 158, 159 
Cotula sect. Strongylosperma (Less.) Benth. 158, 

159 
Cotula abyssinica Schultz-Bip. 158 
Cotula alpina (Hook, f.) Hook. f. 158 
Cotula andreae (E. Phillips) Bremer & Humphries 

158 
Cotula anthemoides L. 158 
Cotula australis (Sieber ex Sprengel) Hook. f. 158 
Cotula barbata DC. 158 
Cotula bipinnata Thunb. 158 
Cotula bracteolata E. Meyer ex DC. 158 
Cotula cabrerae Caro 158 
Cotula ceniifolia DC. 158 
Cotula cinerea del. 158 
Cotula coronopifolia L. 157, 158 
Cotula cotuloides (Steetz) Druce 158 
Cotula cryptocephala Schultz-Bip. ex A. Richards 

158 
Cotula dielsii Muschler 158 
Cotula duckittiae (L. Bolus) Bremer & Humphries 

158 
Cotula eckloniana (DC.) Levyns 158 
Cotula elongata C. B. Vogel 158 
Cotula filifolia Thunb. 158 
Cotula goughensis R. N. R. Brown 157, 158 
Cotula haastii Kirk 159 
Cotula heterocarpa DC. 158 
Cotula hispida (DC.) Harvey 158 
Cotula laxa DC. 158 
Cotula leptalea DC. 158 
Cotula linearifolia Cheeseman 159 
Cotula lineariloba (DC.) Hilliard 158 
Cotula loganii Hutch. 158 
Cotula macroglossa Bolus ex Schltr. 158 
Cotula mariae Bremer & Humphries 158 
Cotula melaleuca Bolus 158 
Cotula membranacea D. Lloyd 159 
Cotula membranifolia Hilliard 158 
Cotula mexicana (DC.) Cabrera 157, 158 
Cotula microglossa (DC.) O. Hoffm. & Kunze ex 

Kunze 158 
Cotula montana Compton 158 
Cotula monticola Simpson 159 
Cotula moseleyi Hemsley 157, 158 
Cotula multifida DC. 159 
Cotula myriophylloides Harvey in Hook. 158 
Cotula nigellifolia (DC.) Bremer & Humphries 158 
Cotula nudicaulis Thunb. 158 
Cotula paludosa Hilliard 158 
Cotula paradoxa Schinz 158 
Cotula pectinata Hook, f . 158 
Cotula pedicellata (Ruiz Lopez & Pavon) Cabrera 

158 



Cotula pedicellata Compton 158, 159 

Cotula pedunculata (Schltr) E. Phillips 159 

Cotula perpusilla Hook. f. 159 

Cotula pterocarpa DC. 159 

Cotula pubescens Desf. 157 

Cotula pusilla Thunb. 159 

Cotula pygmaea Benth. in Benth. & Hook. f. 158 

Cotula radiata O. Hoffm. ex OK. 159 

Cotula radicalis (Killick & Claassen) Hilliard & 

Burtt 159 
Cotula renwickii Cockayne 159 
Cotula rosea Boj. ex Less. 159 
Cotula sericea (Kirk) Cockayne & Allan 159 
Cotula sericea L. f. 159 
Cotula socialis Hilliard 159 
Cotula sororia DC. 159 
Cotula stenophylla K. Koch 159 
Cotula tenella E. Meyer ex DC. 159 
Cotula thunbergii Harvey 159 
Cotula turbinata L. 159 
Cotula umbellata L.f. 160 
Cotula villosa Simpson 159 
Cotula villosa DC. 159 
Cotula vulgaris Levyns 159 
Cotula willcoxii Cheeseman 159 
Cotula zeyheri Fenzl ex Harvey 159 
Cotuleae 75, 76, 83, 

Crossostephium Less. Ill, 113, 117, 120 
Crossostephium artemisioides Less. 120 
Crossostephium chinense (L.) Makino 120 
Cymbopappus B. Nord. 77, 147, 149, 151, 152 
Cymbopappus adenosolen (Harvey) B. Nord. 151, 

152 
Cymbopappus hilliardiae B. Nord. 151 
Cymbopappus lasiopodus (Hutch.) B. Nord. 151 
Cymbopappus piliferus (Thell.) B. Nord. 151 



Daveaua Willk. ex Mariz 147, 149, 153 

Daveaua anthemoides Mariz 153 

Dendranthema (DC.) Des Moul. 77, 106, 110, 111, 

113, 114, 139 
Dendranthema sect. Ajania (Polj.) Kitam. 115 
Dendranthema sect. Arctanthemum Tzvelev 114 
Dendranthema aphrodite (Kitam.) Kitam. 114 
Dendranthema arcticum (L.) Tzvelev 114 
Dendranthema argyrophyllum (Ling) Ling & Shih 

114 
Dendranthema arisanense (Hayata) Ling & Shih 

114 
Dendranthema boreale (Makino) Ling ex Kitam. 

114 
Dendranthema chalchingolicum (Grubov) Bremer 

& Humphries 114 
Dendranthema chanetii (A. Leveille) Shih 114 
Dendranthema coreanum (A. Leveille & Vaniot) 

Vorosch. 114 
Dendranthema crassum (Kitam.) Kitam. 114 
Dendranthema cuneifolium (Kitam.) Bremer & 

Humphries 114 
Dendranthema dichrum Shih 114 
Dendranthema erubescens (Stapf) Tzvelev 114 
Dendranthema glabriusculum (W. Smith) Shih 114 
Dendranthema grandiflorum (Ramat.) Kitam. 114 
Dendranthema hultenii (A. & D. Love) Tzvelev 

114 
Dendranthema hypargyrum (Diels) Ling & Shih 

114 
Dendranthema indicum (L.) Des Moul. 114 
Dendranthema integrifolium (Richardson) Tzvelev 

114 
Dendranthema japonense (Nakai) Kitam. 114 
Dendranthema japonicum (Makino) Kitam. 114 
Dendranthema kurilense Tz\e\c\ 114 
Dendranthema lavanduiifolium (Fischer ex 

Trautv.) Kitam 114 
Dendranthema littorale (Mackawa) Tzvelev 114 
Dendranthema maximowiczii (V. Komarov) 

Tzvelev 114 



Dendranthema miyatojimensc (Kitam.) Hind 114 
Dendranthema mongolicum (Ling) Tzvelev 114 
Dendranthema morifolium (Ramat.) Tzvelev 114 
Dendranthema morii (Hayata) Kitam. 114 
Dendranthema naktongense (Nakai) Tzvelev 114 
Dendranthema nankingcnse (Hand.-Mazz.) Y. R. 

Ling 114 
Dendranthema okiense (Kitam.) Kitam. 114 
Dendranthema oreastrum (Hance) Ling 114 
Dendranthema ornatum (Hemsley) Kitam. 114 
Dendranthema pacificum (Nakai) Kitam. 114, 115 
Dendranthema pallasianum (Fischer ex Besser) 

Vorosch. 114, 115 
Dendranthema parvifolium (Chang) Shih 114 
Dendranthema potentilloides (Hand.-Mazz.) Shih 

114 
Dendranthema rhombifolium Ling & Shih 114 
Dendranthema rupestre (Matsum. ex Koidzumi) 

Kitam. 114 
Dendranthema shiwogiku (Kitam.) Kitam. 114 
Dendranthema sichotense Tzvelev 1 14 
Dendranthema sinchangense (Ueki) Kitam. 114 
Dendranthema sinuatum (Ledeb.) Tzvelev 114 
Dendranthema vestitum (Hemsley) Ling 114 
Dendranthema weyrichii (Maxim.) Tzvelev 114 
Dendranthema xeromorphum Khokr. 114 
Dendranthema yezoense (T. Mack.) Hind 114 
Dendranthema yoshinaganthum (Makino ex 

Kitam.) Kitam. 114 
Dendranthema zawadskii (Herbich) Tzvelev 113 
Dimorphocoma 76, 161 
Diotis Desf. 128 
Diotis candidissima Desf. 128 



Elacanthus F. Muell. 76, 161 
Elachanthemum Ling & Y. R. Ling 116 
Elachanthemum intricatum (Franchet) Ling & Y. 

R. Ling 116 
Endopappus Schultz-Bip. 149, 154, 155 
Ednopappus macrocarpus Schultz-Bip. 154 
Eriocephalus L. 75, 81, 127, 147, 150, 151, 160, 161 
Eriocephalus africanus L. 160, 161 
Eriocephalus aromaticus C. A. Smith 161 
Eriocephalus aspalathoides DC. 161 
Eriocephalus capitellatus DC. 161 
Eriocephalus dinteri S. Moore 161 
Eriocephalus ericoides (L. f.) Druce 161 
Eriocephalus eximius DC. 161 
Eriocephalus glaber Thunb. 161 
Eriocephalus kingesii Merxm. & Eberle 161 
Eriocephalus macroglossus B. Nord. 161 
Eriocephalus microcephalus DC. 161 
Eriocephalus pauperrimus Merxm. & Eberle 161 
Eriocephalus petrophiloides DC. 161 
Eriocephalus pinnatus O. Hoffm. 161 
Eriocephalus pteronioides DC. 161 
Eriocephalus pubescens DC. 161 
Eriocephalus punctulatus DC. 161 
Eriocephalus racemosus L. 161 
Eriocephalus scariosissimus S. Moore 161 
Eriocephalus scariosus DC. 161 
Eriocephalus septulifer DC. 161 
Eriocephalus sericeus Gaudich. ex DC. 161 
Eriocephalus spinescens Burch. 161 
Eriocephalus tenuipes C. A. Smith 161 
Eriocephalus tuberculosus DC. 161 
Eriocephalus umbellulatus Cass. 161 
Eriocephalus xerophilus Schltr 161 
Euanthemideae 75 
Euhelianthees 75 
Euartemisia Gren. & Godron 120 
Eumorphia DC. 77, 92, 94, 95 
Eumorphia corymbosa E. PhilHps 95 
Eumorphia davyi Bolus 95 
Eumorphia dregeana DC. 94, 95 
Eumorphia prostrata Bolus 94, 95 
Eumorphia sericea J. M. Wood & M. Evans 95 
Eumorphia swaziensis Compton 95 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



171 



Eupatorieac 74, 75, 91 

Filifolium Kitam. 112, 116 

Filifolium sibiricum (L.) Kitam. 116 

Formania W. W. Smith 161 

Foveolina Kallersjo 77, 147, 149, 151, 154, 155, 157 

Foveolina albida (DC.) Kallersjo 155 

Foveolina albidiformis (Thell.) Kallersjo 155 

Foveolina dichotoma (DC.) Kallersjo 155 

Foveolina schinziana (Thell.) Kallersjo 155 

Foveolina tenella (DC.) Kallersjo 155 

Glossanthis Polj. 97 
Glossopappus Kunze 139, 143, 144 
Glossopappus chrysanthemoides Kunze 143 
Glossopappus macrotus (Durieu) Briq. in Burnat 

143 
Gnaphalieae 161 
Gonosperminae Bremer & Humphries 74, 76, 77, 

106, 107, 145 
Gonospermum Less. 76, 77, 83, 107-108 
Gonospermum canariense (DC.) Less. 108 
Gonospermum elegans (Cass.) DC. 108 
Gonospermum fruticosum (C. Smith ex Link) 

Less. 106-108 
Gonospermum gomerae Bolle 108 
GymnocUne Cass. 100 

Gymnopentzia Bcnth. in Benth. & Hook. f. 92, 95 
Gymnopentzia bifurcata Benth. 95 
Gymnostyles Juss. 159, 160 
Gymnostyles stolonifera (Brot.) Tutin 160 

Handelia Hcimerl 108, 109, 110 

Handelia trichophylla (Schrenk) Heimerl 108, 110 

Handeliinae Bremer & Humphries 74, 76, 77, 108, 

109 
Helenieae 81, 
Heleniees 75 

Heliantheae Cass. 74, 75, 77, 80-82, 90, 91 
Helianthees 75 

Heliocauta Humphries 82, 99, 100, 106 
Heliocauta atlantica (Litard & Maire) Humphries 

106 
Hemipappus K. Koch 100, 101 
Hemipappus argenteus (Lam.) Tzvelev 101 
Hemipappus canus K. Koch 101 
Heteranthemis Schott 134, 135, 136 
Heteranthemis viscidehirta Schott 135 
Heteromera Pomel 147, 149, 153 
Hcteromera fuscata (Desf.) Pomel 153 
Heteromera philaenorum Maire & Weller 153 
Hilliardia B. Nord. 77, 147, 150, 157 
Hilliardia zuurbergensis (Oliver) B. Nord. 157 
HippiaL. 147, 150, 151, 160 
Hippia bolusae Hutch. 160 
Hippia frutescens (L.) L. 160 
Hippia hirsuta DC. 160 
Hippia hutchinsonii Merxm. 160 
Hippia integrifolia Less. 160 
Hippia montana Compton 160 
Hippia pilosa (P. Bergius) Druce 160 
Hippia trilobata Hutch. 160 
Hippolytia Polj. 100, 105, 106, 145 
Hippolytia alashanensis (Ling) Shih 106 
Hippolytia crassicollum (Rech. fil.) Bremer & 

Humphries 106 
Hippolytia darvasica (Winkler) Polj. 105, 106 
Hippolytia delavayi (W. Smith) Shih 106 
Hippolytia desmantha Shih 106 
Hippolytia dolicliophylla (Kitam.) Bremer & 

Humphries 106 
Hippolytia glomerata Shih 106 
Hippolytia gossypina (C. B. Clarke) Shih 106 
Hippolytia herderi (Regel & Schmalh.) Polj. 106 
Hippolytia kaschgarica (H. Kraschen.) Polj. 106 
Hippolytia kennedyi (Dunn) Ling 106 
Hippolytia longifolia (Wallich) Shih 106 
Hippolytia megacephala (Rupr.) Polj. 106 



Hippolytia nana (C. B. Clarke) Shih 106 
Hippolytia schugnanica (Winkler) Polj. 106 
Hippolytia senecionis (Besser) Polj. 106 
Hippolytia syncalathiformis Shih 106 
Hippolytia tomentosa (DC.) Tzvelev 106 
Hippolytia trifida (Turcz.) Polj. 106 
Hippolytia yunnanensis (Jeffrey) Shih 106 
Hulteniella Tz\e\e\ 114 

Hultenietla imegrifolium (Richardson) Tzvelev 114 
Hymenolepis Cass. 77, 91, 92, 95 
Hymenolepis cynopus Bremer •& Kallersjo 95 
Hymenolepis dentata (DC.) Kallersjo 95 
Hymenolepis gnidioides (S. Moore) Kallersjo 95 
Hymenolepis incisa DC. 95 
Hymenolepis indivisa (Harvey) Kallersjo 95 
Hymenolepis parviflora (L.) DC. 95 
Hymenolepis speciosa (Hutch.) Kallersjo 95 
Hymenopappus V Herit 161 
Hymenostemma (Kunze) Willk. 77, 136, 137, 138, 

140, 141 
Hymenostemma paludosum (Poiret) Pomel 142 
Hymenostemma pseudanthemis (Kunze) Willk. 

141 

Inezia E. Phillips 144, 145, 146 

Inezia integrifolia (Klatt) E. Phillips 146 

Inezia speciosa Brusse 146 

Inulanthera Kallersjo 76, 77, 83, 91, 107, 108, 145 

Inulanthera brownii (Hochr.) Kallersjo 108 

Inulanthera calva (Hutch.) Kallersjo 108 

Inulanthera coronopifolia (Harvey) Kallersjo 108 

Inulanthera dregeana (DC.) Kallersjo 108 

Inulanthera leucoclada (DC.) Kallersjo 108 

Inulanthera montana (J. M. Wood & M. Evans) 

Kallersjo 108 
Inulanthera nuda Kallersjo 108 
Inulanthera schistostephioides (Hiern) Kallersjo 

108 
Inulanthera thodei (Bolus) Kallersjo 108 
Inulanthera tridens (Oliver) Kallersjo 108 
Inuleae 75 

Ischnea F. Muell. 161 
Ismelia Cass. 77, 135, 136 
Ismelia carinata (Schousboe) Schultz-Bip. 135 
Ismelia versicolor Cass. 135 
Isoetopsis Turcz. 76, 161 

Kaschgaria Polj. 113, 117, 121 

Kaschgaria brachanthemoides (Winkler) Polj. 117 

Kaschgaria komarovii (H. Kraschen. & N. 

Rubtzov) Polj. 117 
Kremeria myconis (L.) Maire 143 
Kremeria paludosa Durieu 143 

Lagenophora Cass. 161 
Lasiospermum Lagasca 75, 83, 91, 92, 94 
Lasiospermum bipinnatum (Thunb.) Druce 94 
Lasiospermum brachyglossum DC. 94 
Lasiospermum erectum (Lam.) Druce 94 
Lasiospermum pedunculare Lagasca 94 
Lasiospermum poterioides Hutch. 94 
Lasiospermum radiatum Trevir. 94 
Lepidolopha Winkler 100, 101, 105 
Lepidolopha fedtschenkoana Knorr. 105 
Lepidolopha filifolia Pavlov 105 
Lepidolopha gomohtzkii Kovalevsk. & Safralieva 

105 
Lepidolopha karatavica Pavlov 105 
Lepidolopha komarovii Winkler 105 
Lepidolopha krascheninnikovii Kovalevsk. & 

Safralieva 105 
Lepidolopha mogoltavica (H. Kraschen.) H. 

Kraschen. 105 
Lepidolopha nuratavica H. Kraschen. 105 
Lepidolopha talasica Kovalevsk. & Safralieva 105 
Lepidolopsis Polj. 104, 108, 109 
Lepidolopsis turkestanica (Regel & Schmalh.) 

Polj. 109 



Lepidophorum Necker ex Cass. 77, 136, 137, 139, 

145 
Lepidophorum repandum (L.) DC. 139 
Lepidostephium Oliver 161 
Leptinella Cass. 76, 83, 147, 148, 150, 159 
Leptinella albida (D. Lloyd) D. Lloyd & C. Webb 

159 
Leptinella altilitoralis (P. Royen & D. Lloyd) D. 

Lloyd & C. Webb 159 
Leptinella atrata (Hook, f.) D. Lloyd & C. Webb 

159 
Leptinella calcarea (D. Lloyd) D. Lloyd & C. 

Webb 159 
Leptinella dendyi (Cockayne) D. Lloyd & C. 

Webb 159 
Leptinella dioica Hook. f. 159 
Leptinella dispersa (D. Lloyd) D. Lloyd & C. 

Webb 159 
Leptinella drummondii (Benth.) D. Lloyd & C. 

Webb 159 
Leptinella featherstonii F. Muell. 159 
Leptinella filicula (Hook, f.) Hook. f. 159 
Leptinella filiformis (Hook, f.) D. Lloyd & C. 

Webb 159 
Leptinella goyenii (Petrie) D. Lloyd & C. Webb 

159 
Leptinella intermedia (D. Lloyd) D. Lloyd & C. 

Webb 159 
Leptinella lanata Hook. f. 159 
Leptinella leptoloba (Mattf.) D. Lloyd & C. Webb 

159 
Leptinella longipes Hook. f. 159 
Leptinella maniototo (Petrie) D. Lloyd & C. Webb 

159 
Leptinella minor Hook. f. 159 
Leptinella nana (D. Lloyd) D. Lloyd & C. Webb 

159 
Leptinella pectinata (Hook, f.) D. Lloyd & C. 

Webb 159 
Leptinella plumosa Hook. f. 159 
Leptinella potentilhna (F. Muell.) Druce 159 
Leptinella pusilla Hook. f. 159 
Leptinella pyrethrifolia (Hook, f.) D. Lloyd & C. 

Webb 159 
Leptinella reptans (Benth.) D. Lloyd & C. Webb 

159 
Leptinella rotundata (Cheeseman) D. Lloyd & C. 

Webb 159 
Leptinella sarawaketensis (P. Royen & D. Lloyd) 

D. Lloyd & C. Webb 159 
Leptinella scariosa (Cass.) Franchet 159 
Leptinella serrulata (D. Lloyd) D. Lloyd & C. 

Webb 159 
Leptinella squalida Hook. f. 159 
Leptinella tenella (Cunn.) D. Lloyd & C. Webb 

159 
Leptinella traillii (Kirk) D. Lloyd & C. Webb 159 
Leptinella wilhelminensis (P. Royen) D. Lloyd & 

C. Webb 159 
Leucampyx A. Gray 161 
Leucanthemella Tzvelev 77, 138, 139, 140 
Leucanthemella linearis (Matsum.) Tzvelev 139, 

140 
Leucanthemella serotina (L.) Tzvelev 137, 139, 140 
Leucantheminae Bremer & Humphries 
74, 77, 81, 82, 83, HI, 128, 131, 136-139, 140, 145, 

149 
Leucanthemopsis (Giroux) Heyw. 77, 136, 137, 

138, 140, 141 
Leucanthemopsis alpina (L.) Heyw. 140 
Leucanthemopsis flaveola (Hoffsgg & Link) Heyw. 

140 
Leucanthemopsis longipectinata (Font Ouer) 

Heyw. 140 
Leucanthemopsis minima (Villars) Marchi 140 
Leucanthemopsis pallida (Miller) Heyw. 140 
Leucanthemopsis f)ectinata (L.) Lopez Gonzalez & 
Jarvis 140 



172 



K. BREMER AND C. J. HUMPHRIES 



Leucanthcmopsis pulverulenta (Lagasca) Heyw. 

140 
Leucanthemopsis radicans (Cav.) Heyw. 140 
Leucanthemopsis tatrae (Vierh.) Holub 140 
Leucanthemopsis tomentosa (Lois.) Marchi 140 
Leucanthemopsis trifurcata (Desf.) Alavi 140, 143 
Leucanthemum Miller 77, 82, 83, 137, 138, 139, 

141, 142 
Leucanthemum subgen. Chrysanthemopsis Maire 

141 
Leucanthemum sect. Rhodanthemum 141 
Leucanthemum adustum (Koch) Gremli 141 
Leucanthemum aligulatum Vogt 141 
Leucanthemum arundanum (Boiss.) Cuatrec. 141, 

142 
Leucanthemum atlanticum (Ball) Maire 142 
Leucanthemum atratum (Jacq.) DC. 141 
Leucanthemum briquetii Maire 142 
Leucanthemum burnatii Briq. & Cav. 141 
Leucanthemum catalaunicum Vogt 141 
Leucanthemum catananche (Ball) Maire 142 
Leucanthemum chloroticum A. Kerner & Murb. 

141 
Leucanthemum corsicum (Less.) DC. 141 
Leucanthemum crassifolium (Lange) Willk. in 

Willk. & Lange 141 
Leucanthemum cuneifolium Le Grand ex Coste 

141 
Leucanthemum decipiens Pomel 142 
Leucanthemum delarbrei Timb.-Lagr. 141 
Leucanthemum depressum (Ball) Maire 142 
Leucanthemum discoideum (All.) Coste 141, 144 
Leucanthemum favargeri Vogt 141 
Leucanthemum fontanesii Boiss. & Reuter 141 
Leucanthemum gayanum (Cosson & Durieu) Maire 

142 
Leucanthemum gaudinii Dalla Torre 141 
Leucanthemum gracilicaule (Duf.) Alavi & Heyw. 

141 
Leucanthemum graminifolium (L.) Lam. 141 
Leucanthemum heterophyllum (Willd.) DC. 141 
Leucanthemum hosmariense (Ball) Font Quer 142 
Leucanthemum ircutianum DC. 141 
Leucanthemum laciniatum Huter, Porta & Rigo 

141 
Leucanthemum lacustre (Brot.) Samp. 141 
Leucanthemum leucolepis (Briq. & Cav.) Horvatic 

141 
Leucanthemum maestracense Vogt & Hellwig 141 
Leucanthemum mairei Humbert 142 
Leucanthemum maresii (Cosson) Maire 142 
Leucanthemum maroccanum (Battand.) Maire 142 
Leucanthemum maximum (Ram.) DC. 141 
Leucanthemum meridionale Le Grand 141 
Leucanthemum merinoi Vogt & Castroviejo 141 
Leucanthemum mesatlanticum Emb. & Maire 142 
Leucanthemum monspeliense (L.) Coste 141 
Leucanthemum montserratianum Vogt 141 
Leucanthemum pallens (Gay) DC. 141 
Leucanthemum paludosum (Poiret) Bonnet & 

Barratte 141, 142 
Leucanthemum praecox (Horvatic) Horvatic 141 
Leucanthemum pluriflorum Paul. 141 
Leucanthemum pseudo-catananche Maire 142 
Leucanthemum reboudianum Pomel 142 
Leucanthemum redieri Maire 142 
Leucanthemum setabense DC. 142 
Leucanthemum subglaucum De Laramb. 141 
Leucanthemum sylvaticum (Hoffsgg & Link) 

Nyman 141 
Leucanthemum vulgare Lam. 136, 141 
Leucanthemum waldensteinii (Schultz-Bip.) 

Pouzar 141 
Leucocyclus Boiss. 77, 126, 127, 130 
Leucocyclus formosus Boiss. 130 
Leucoglossum Wilcox, Bremer & Humphries 77, 

139, 140, 141, 142, 143 



Leucoglossum decipiens (Pomel) Wilcox, Bremer & 

Humphries 142 
Leucoglossum paludosum (Poiret) Wilcox, Bremer 

& Humphries 141, 142 
Leucoglossum reboudianum (Pomel) Wilcox, 

Bremer & Humphries 142 
Leucoptera B. Nord. 147, 150, 157 
Leucoptera nodosa (Thunb.) B. Nord. 157 
Leucoptera oppositifolia B. Nord. 157 
Leucoptera subcarnosa B. Nord. 157 
Ley sera L. 161 

Lidbeckia P. Bergius 144, 145, 146 
Lidbeckia lobata Thunb. 146 
Lidbeckia pectinata P. Bergius 145, 146 
Lidbeckia quinqueloba (L. f.) Cass. 146 
Lidbeckieae 75 
Lonas Adans. 149, 153, 154 
Lonas annua (L.) Vines & Druce 154 
Lugoa DC. 76, 83, 107 
Lugoa revoluta (C. Smith ex Link) DC. 107 



Marasmodes DC. 147, 149, 151, 152 
Marasmodes adenosolen Harvey 152 
Marasmodes dummeri Bolus ex Hutch. 152 
Marasmodes oligocephalus DC. 152 
Marasmodes polycephalus DC. 152 
Marasmodes undulata Compton 152 
Matricaria L. 75, 77, 83, 146, 147-149, 153, 154, 

155, 156, 158 
Matricaria andreae E. Phillips 158 
Matricaria armeniaca Rauschert 156 
Matricaria aserbaidshanica Rauschert 156 
Matricaria aurea (Loefl.) Schultz-Bip. 154 
Matricaria auriculata (Boiss.) Ball 156 
Matricaria australis (Pobed.) Rauschert 156 
Matricaria breviradiata (Ledeb.) Rauschert 156 
Matricaria capensis L. 153 
Matricaria caucasica (Willd.) Poiret 156 
Matricaria chamomilla L. 153 
Matricaria chamomilla auct. non L. 154 
Matricaria colchica (Manden.) Rauschert 156 
Matricaria conoclinia (Boiss. & Balansa) Nyman 

156 
Matricaria corymbosa Desr. in Lam. 156 
Matricaria daghestanica (Rupr. ex Boiss.) 

Rauschert 156 
Matricaria decipiens (Fischer & C. Meyer) K. Koch 

156 
Matricaria disciformis (C. Meyer) DC. 156 
Matricaria discoidea DC. 154 
Matricaria elongata (Fischer & C. Meyer ex DC.) 

Hand.-Mazz. 156 
Matricaria froedinii (Rcch. f.) Rauschert 156 
Matricaria globifera (Thunb.) Fenzl ex Harvey 153 
Matricaria grossheimii (Fed.) Rauschert 156 
Matricaria halepensis Rauschert 156 
Matricaria hirta (Thunb.) DC. 153 
Matricaria hygrophila (Bornm.) Rauschert 156 
Matricaria inodora L. 153, 154, 156 
Matricaria karjaginii (Manden & Sof.) Rauschert 

156 
Matricaria lasiocarpa Boiss. 154 
Matricaria ledebourii (Schultz-Bip.) Schischk. 154 
Matricaria lesbiaca (Candargy) Rauschert 156 
Matricaria limosa (Maxim.) Kudo 156 
Matricaria macrotis Rech. f. 154 
Matricaria maritima L. 156 
Matricaria matricarioides Porter ex Britton 154 
Matricaria microcephala K. Koch 156 
Matricaria monticola (Boiss. & Huet) Rauschert 

156 
Matricaria multiflora Fenzl ex Harvey 153 
Matricaria nigellifolia DC. 158 
Matricaria occidentalis (Greene) Rauschert 154 
Matricaria oreades Boiss. 156 
Matricaria parviflora (Willd.) Poiret 156 
Matricaria perforata Merat 154, 156 
Matricaria pichleri (Boiss.) Rauschert 156 



Matricaria pubescens (Dcsf.) Schultz-Bip. 157 
Matricaria recutita L. 146, 153, 154 
Matricaria rosella (Boiss. & Orph.) Nyman 156 
Matricaria rupestris (Sommier & Levier) Rauschert 

156 
Matricaria sevanesis (y^dtnd^n.) Rauschert 156 
Matricaria songarica Bunge 154 
Matricaria sub nivalis {Vohcd.) Rauschert 156 
Matricaria szowitzii (DC.) Rauschert 156 
Matricaria tchihatchewii (Boiss.) Voss 156 
Matricaria tempskyana (Freyn & Sint.) Rauschert 

156 
Matricaria tetragonosperma (Schum.) Hara & 

Kitam. 156 
Matricaria transcaucasica (Manden.) Rauschert 156 
Matricaria trichophylla (Boiss.) Boiss. 156 
Matricaria tzvelevii Pobed. 154, 156 
Matricariinae Bremer & Humphries 
74, 75-77, 83, 84, 127, 128, 145, 146-151, 155 
Mausolea Polj. 82, 111, 121, 125 
Mausolea eriocarpa (Bunge) Polj. 125 
Mecomischus Cosson ex Benth. in Benth. & Hook. 

f. 77, 126, 127, 130 
Mecomischus geslini (Cosson) Cosson 130 
Mecomischus halimifolius (Munby) Hochr. 130 
Mecomischus pedunculatus (Cosson & Durieu) 

Maire 130 
Microcephala Pobed. 77, 147, 149, 154 
Microcephala afghanica Podl. 154 
Microcephala deserticola Podl. 154 
Microcephala lamellata (Bunge) Pobed. 154 
Microcephala lasiocarpa (Boiss.) Pobed. 154 
Microcephala subglobosa (H. Kraschen.) Pobed. 

154 
Microcephala turcomanica (Winkler) Pobed. 154 
Myxopappus Kallersjo 77, 90, 147, 149, 151, 155, 

157 
Myxopappus acutilobus (DC.) Kallersjo 155 
Myxopappus hereroensis (O. Hoffm.) Kallersjo 

155 

Nananthea DC. 76, 77, 131, 132, 134 
Nananthea perpusilla (Lois.) DC. 134 
Nassauvineae 75 
NeopallasiaPolj.81, 121, 125 
Neopallasia pectinata (Pallas) Polj. 125 
Neopallasia tibetica Y. R. Ling 125 
Neopallasia yunnancnsis (Pampan.) Y. R. Ling 125 
Nipponanthemum Kitam. 77, 137, 139 
Nipponanthemum nipponicum (Franchet ex 

Maxim.) Kitam. 139 
Nivellea Wilcox, Bremer & Humphries 77, 137, 

138, 140 
Nivellea nivellei (Braun-Blanquet & Maire) Wilcox, 

Bremer & Humphries 140 

Oedera L. 161 

Oligosporus Cass. 120, 121 

Oncosiphon Kallersjo 77, 82, 147-151, 152, 153, 

157 
Oncosiphon africanum (P. Bergius) Kallersjo 153 
Oncosiphon glabratum (Thunb.) Kallersjo 153 
Oncosiphon grandiflorum (Thunb.) Kallersjo 153 
Oncosiphon intermedium (Hutch.) Kallersjo 153 
Oncosiphon piluliferum (L. f.) Kallersjo 152, 153 
Oncosiphon sabulosum (Wolley-Dod) Kallersjo 

153 
Oncosiphon schlechteri (Bolus) Kallersjo 153 
Oncosiphon suffruticosum (L.) Kallersjo 153 
Opisthopappus Shih 99, 104 
Opisthopappus longilobus Shih 104 
Opisthopappus taihangensis (Ling) Shih 104 
Ormenis (Cass.) Cass. 130 
Ormenis lonadioides (Cosson) Maire 131 
Ormenis mixta (L.) Dumort. 130 
Ormenis nobilis (L.) Gay 130 
Ormenis praecox (Link) Briq. & Cavill. 130 
Ormenis santolinoides (Munby) Harling 130 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



173 



Osmitopsis Cass. 144, 145 

Osmitopsis afra (L.) Bremer 145 

Osmitopsis asteriscoides (P. Bergius) Less. 145 

Osmitopsis dentata (Thunb.) Bremer 145 

Osmitopsis glabra Bremer 145 

Osmitopsis nana Schltr 145 

Osmitopsis osmitoides (Less.) Bremer 145 

Osmitopsis parvifolia (DC.) Hofmeyer 145 

Osmitopsis pinnatifida (DC.) Bremer 145 

Osmitopsis tenuis Bremer 145 

Otanthus Hoffsgg & Link 82, 126, 127, 128 

Otanthus maritimus (L.) Hoffsgg & Link 128 

Otochlamys DC. 157 

Otochlamys eckloniana DC. 158 

Otochlamys pedunculata Schltr 159 

Otospermum Willk. 149, 153 

Otospermum glabrum (Lagasca) Willk. 153 



Pentzia Thunb. 76, 77, 147-149, 151, 152, 154 

Pentzia annua DC. 155 

Pentzia argentea Hutch. 151 

Pentzia bolusii Hutch. 151 

Pentzia calcarea Kies 151 

Pentzia calva S. Moore 152 

Pentzia cooperi Harvey 152 

Pentzia crenata Thunb. 151 

Pentzia dentata (L.) OK. 151, 152 

Pentzia eenii S. Moore 152 

Pentzia elegans DC. 152 

Pentzia galpinii Hutch. 155 

Pentzia globifera (Thunb.) Hutch. 153 

Pentzia globosa Less. 152 

Pentzia hesperidum Maire & Wilczek 152 

Pentzia incana (Thunb.) OK. 152 

Pentzia lanata Hutch. 152 

Pentzia membranacea Hutch. 155 

Pentzia laxa Bremek. & Oberm. 152 

Pentzia monocephala S. Moore 152 

Pentzia monodiana Maire & Wilczek 151, 152 

Pentzia nana Burch. 152 

Pentzia peduncularis B. Nord. 152 

Pentzia pinnatisecta Hutch. 152 

Pentzia punctata Harvey 152 

Pentzia quinquefida (Thunb.) Less. 152 

Pentzia schistostephioides M. Taylor 108 

Pentzia sphaerocephala DC. 152 

Pentzia spinescens Less. 152 

Pentzia tanacetifolia (L.) Hutch. 153 

Pentzia tomentosa B. Nord. 152 

Pentzia tortuosa (DC.) Fenzl ex Harvey 152 

Pentzia viridis Kies 152 

Peyrousea DC. 160 

Peyrousea umbellata (L. f.) Fourc. 160 

Phaeocephalus S. Moore 95 

Phaeocephalus gnidioides S. Moore 95 

Phaeostigma Muld. 82, 110, 111, 112, 115, 116 

Phaeostigma quercifolium (W. Smith) Muld. 116 

Phaeostigma salicifolium (Mattf.) Muld. 115, 116 

Phaeostigma variifolium (Chang) Muld. 116 

Phalacrocarpum (DC.) Willk. 77, 137, 138, 140 

Phalacrocarpum anomalum Cout. 140 

Phalacrocarpum hoffmannseggii (Samp.) Lainz 140 

Phalacrocarpum oppositifolium (Brot.) Willk. 140 

Phymaspermum Less. 77, 91, 92, 94, 95 

Phymaspermum acerosum (DC.) Kallersjo 94 

Phymaspermum aciculare (E. Meyer ex Harvey) 

Benth. ex B. D. Jackson 94 
Phymaspermum appressum Bolus 94 
Phymaspermum argcnteum Brusse 94 
Phymaspermum athanasioides (S. Moore) 

Kallersjo 94 
Phymaspermum bolusii (Hutch.) Kallersjo 94 
Phymaspermum equisetoides Thell. 94 
Phymaspermum erubescens (Hutch.) Kallersjo 94 
Phymaspermum junceum Less. 94 
Phymaspermum leptophyllum (DC.) Benth. ex B. 

D. Jackson 94 
Phymaspermum montanum (Hutch.) Kallersjo 94 



Phymaspermum parvifolium (DC.) Benth. ex B. 

D. Jackson 94 
Phymaspermum peglerae (Hutch.) Kallersjo 94 
Phymaspermum pinnatifidum (Oliver) Kallersjo 94 
Phymaspermum schroteri Compton 94 
Phymaspermum scoparium (DC.) Kallersjo 94 
Phymaspermum villosum (Hilliard) Kallersjo 94 
Phymaspermum woodii (Thell.) Kallersjo 94 
Picrothamnus Nutt. 111-113, 121, 125, 126 
Picrothamnus desertorum Nutt. 125 
Plagiocheilus Am. ex D.C. 76, 161 
Plagius L'Herit. ex DC. 136, 138, 139, 143, 144 
Plagius ageratifolius (Desf.) L'H6rit. ex DC. 143 
Plagius flosculosus (L.) Alavi & Heyw. 143, 144 
Plagius grandis (L.) Alavi & Heyw. 144 
Polychrysum (Tzvelev) Kovalevsk. 108, 109 
Polychrysum tadshikorum (Kudr.) Kovalevsk. 109 
Prolongoa Boiss. 77, 82, 83, 136, 137, 138, 140, 141 
Prolongoa sect. Hymenostemma Kunze 140, 141, 

142 
Prolongoa hispanica Lopez Gonzalez & Jarvis 141 
Prolongoa pectinata auci. 141 
Pseudocadiscus Lisowski 161 
PseudohandeHa Tzvelev 108, 109, 110 
Pseudohandeha umbellifera (Boiss.) Tzvelev 109 
Psilostrophe D.C. 161 
Pyrethrum Zinn 98, 100, 101, 105, 106 
Pyrethrum sect. Dendranthema DC. 113 
Pyrethrum sect. Leptanthemum Tzvelev 104 
Pyrethrum sect. Leucanthemopsis (Giroux) Tzvelev 

101 
Pyrethrum sect. Richteria (Karelin & Kir.) Tzvelev 

101 
Pyrethrum sect. Trichanthemopsis Tzvelev 101, 105 
Pyrethrum sect. Xylopyrethrum Tzvelev 101 
Pyrethrum abrotanifolium Bunge ex Ledeb. 101 
Pyrethrum alatavicum (Herder) O. & B. Fedtsch. 

101 
Pyrethrum arassanicum (Winkler) O. & B. 

Fedtsch. 98 
Pyrethrum arctodzhungaricum Golosk. 101 
Pyrethrum aromaticum Tzvelev 101 
Pyrethrum arundanum Boiss. 142 
Pyrethrum atkinsonii (C. B. Clarke) Ling & Shih 

101 
Pyrethrum aucheranum DC. 101 
Pyrethrum balsamita (L.) Willd. 102 
Pyrethrum balsamitoides (Nab.) Tzvelev 102 
Pyrethrum chamaemelifolium (Sommier & Levier) 

Sosn. 102 
Pyrethrum changaicum H. Kraschen. ex Grubov 

102 
Pyrethrum cinerariifolium Trevir 102 
Pyrethrum clusii Fischer ex Reichb. 102 
Pyrethrum coccineum (Willd.) Vorosch. 102 
Pyrethrum corymbiforme Tzvelev 102 
Pyrethrum corymbosum (L.) Willd. 102 
Pyrethrum daghestanicum (Rupr. ex Boiss.) Rupr. 

ex Flerov 102 
Pyrethrum demelrii Manden. 102 
Pyrethrum deserticola Murb. 143 
Pyrethrum discoideum Ledeb. 154 
Pyrethrum djilgense (Franchet) Tzvelev 98 
Pyrethrum dolomiticum Galushko 102 
Pyrethrum fruticulosum Biehler 103 
Pyrethrum galae Popov 102 
Pyrethrum galushkoi Prima 102 
Pyrethrum gayanum Cosson & Durieu 142 
Pyrethrum glanduliferum Sommier & Levier 102 
Pyrethrum grossheimii Sosn. 102 
Pyrethrum heldreichianum Fenzl ex Tchich. 101 
Pyrethrum hissaricum H. Kraschen. 102 
Pyrethrum karelinii H. Kraschen. 102 
Pyrethrum kaschgaricum H. Kraschen. 102 
Pyrethrum kelleri (Krylov & Plotn.) H. Kraschen. 

'l02 
Pyrethrum komarovii Sosn. 103 
Pyrethrum kotschyi Boiss. 102 



Pyrethrum krylovianum H. Kraschen. 102 

Pyrethrum kubense Grossh. 102 

Pyrethrum lanuginosum (Schultz-Bip. & Herder) 

Tzvelev 102 
Pyrethrum leptophyllum Steven 102 
Pyrethrum lingulatum Boiss. 105 
Pyrethrum macrocarpum (Cosson & Kralik) Alavi 

154 
Pyrethrum macrophyllum (Waldst. & Kit.) Willd. 

102 
Pyrethrum majus (Desf.) Tzvelev 102 
Pyrethrum maresii Cosson 142 
Pyrethrum marionii Albov 102 
Pyrethrum mikeschinii Tzvelev 103 
Pyrethrum neglectum Tzvelev 98 
Pyrethrum ordubadense Manden. 103 
Pyrethrum oxylepis (Bordz.) Tzvelev 103 
Pyrethrum parthenifolium Willd. 103 
Pyrethrum parthenifolium var. peucedanifolia Sosn. 

103 
Pyrethrum parthenium (L.) Smith 103 
Pyrethrum petrareum Shih 103 
Pyrethrum peucedanifolium (Sosn.) Manden. 103 
Pyrethrum poteriifolium Ledeb. 103 
Pyrethrum pulchellum Turcz. 103 
Pyrethrum pulchrum Ledeb. 103 
Pyrethrum punctatum (Desr.) Bordz. ex Sosn. 103 
Pyrethrum pyrethroides (Karelin & Kir.) B. 

Fedtsch. & H. Kraschen. 98 
Pyrethrum richterioides (Winkler) H. Kraschen. 

103 
Pyrethrum roseum (Adams) M. Bieb. 103 
Pyrethrum semenovii (Herder) Winkler ex O. & B. 

Fedtsch. 103 
Pyrethrum sericeum (Adams) M. Bieb. 

103Pyrethrum sevanense Sosn. ex Grossh. 103 
Pyrethrum silaifolium Steven 103 
Pyrethrum santolinoides DC. 103 
Pyrethrum songaricum Tzvelev 103 
Pyrethrum sorbifolium Boiss. 103 
Pyrethrum tatsienense (Bureau & Franchet) Ling ex 

Shih 103 
Pyrethrum tianshanicum H. Kraschen. 97, 101, 105 
Pyrethrum transiliense (Herder) Regel & Schmalh. 

98 
Pyrethrum tricholobum Sosn. ex Manden. 103 
Pyrethrum trichophyllum Sosn. 103 
Pyrethrum yabrudae Mout. 104 



Relhania V Herit. emend. Anderb. & Bremer 161 
Rennera Merxm. 77, 147-149, 152 
Rennera eenii (S. Moore) Kallersjo 152 
Rennera laxa (Bremek. & Oberm.) Kallersjo 152 
Rennera limnophila Merxm. 152 
Rhetinolepis Cosson 126, 130, 131 
Rhetinolepis lonadioides Cosson 130, 131 
Rhodanthemum (Vogt) Bremer & Humphries 138, 

141, 142 
Rhodanthemum arundanum (Boiss.) Wilcox, 

Bremer & Humphries 141, 142 
Rhodanthemum atlanticum (Ball) Wilcox, Bremer 

& Humphries 142 
Rhodanthemum briquetii (Maire) Wilcox, Bremer 

& Humphries 142 
Rhodanthemum catananche (Ball) Wilcox, Bremer 

& Humphries 142 
Rhodanthemum depressum (Ball) Wilcox, Bremer 

& Humphries 142 
Rhodanthemum gayanum (Cosson & Durieu) 

Wilcox, Bremer & Humphries 142 
Rhodanthemum hosmariense (Ball) Wilcox, 

Bremer & Humphries 142 
Rhodanthemum maresii (Cosson) Wilcox, Bremer 

& Humphries 142 
Rhodanthemum maroccanum (Battand.) Wilcox, 

Bremer & Humphries 142 
Rhodanthemum mesatlanticum (Emb. & Maire) 

Wilcox, Bremer & Humphries 142 



174 



K. BREMER AND C. J. HUMPHRIES 



Rhodanthemum pseudo-catananche (Maire) 

Wilcox, Bremer & Humphries 142 
Rhodanthemum redieri (Maire) Wilcox, Bremer & 

Humphries 142 
Richteria Karelin & Kir. 77, 96, 98, 105 
Richteria djilgense (Franchet) Bremer & 

Humphries 98 
Richteria leontopodium Winkler 98 
Richteria pyrethroides Karelin & Kir. 98 



Santolina L. 77, 126, 127, 128 

Santolina annua L. 155 

Santohna chamaecyparissus L. 128 

SantoHna elegans Boiss. ex DC. 128 

Santolina insularis (Gennari ex Fiori) Arrig. 128 

Santolina hgustica Arrig. 128 

Santolina marchi Arrig. 128 

Santolina oblongifolia Boiss. 128 

Santolina rosmarinifolia L. 128 

Santolina sinaica Fresen. 103 

Santolina viscosa Lagasca 128 

Santohneae 75 

Schistostephium Less. 147, 151, 160 

Schistostephium crataegifolium (DC.) Fenzl ex 

Harvey 160 
Schistostephium dactyliferum Hutch. 160 
Schistostephium flabelhforme Less. 160 
Schistostephium griseum (Harvey) Hutch. 160 
Schistostephium heptalobum (DC.) Oliver & Hiern 

160 
Schistostephium hippiifoHum (DC.) Hutch. 160 
Schistostephium mollissimum Hutch. 160 
Schistostephium oxylobum S. Moore 160 
Schistostephium rogersii Hutch. 160 
Schistostephium rotundifolium (DC.) Fenzl ex 

Harvey 160 
Schistostephium saxicola Hutch. 160 
Schistostephium scandens Hutch. 160 
Schistostephium umbellatum (L. f.) Bremer & 

Humphries 160 
Schistostephium villosum Hutch. 160 
Sclerorhachis (Rech. f.) Rech. f. 76, 82, 106, 108, 

109, 110 
Sclerorhachis caulescens (Aitch. & Hemsley) 

Rech. f. 110 
Sclerorhachis leptoclada Rech.f 110 
Sclerorhachis platyrachis (Boiss.) Podl. ex Rech. f. 

110 
Sclerorhachis polysphaera Rech. f. 110 
Senecioneae 74, 75, 161 
Senecionideae 75 
Senecionees 75 
Seriphidium (Besser ex Hook.) Fourr. 77, 111-113, 

116,117-120 
Seriphidium algeriense (Filat.) Y. R. Ling 118 
Seriphidium amoenum (Polj.) Polj. 118 
Seriphidium aralense (H. Kraschen.) Polj. 118 
Seriphidium arbusculum (Nutt.) W. A. Weber 118 
Seriphidium arenicolum (H. Kraschen. ex Polj.) Y. 

R. Ling 118 
Seriphidium argilosum (Beetle) Bremer & 

Humphries 118 
Seriphidium assurgens (Filat.) Bremer & 

Humphries in Y. R. Ling 118 
Seriphidium aucheri (Boiss.) Ling & Y. R. Ling 

118 
Seriphidium badhysi (Krasch. & Lincz. ex Polj.) 

Polj. C. Asia. 118 
Seriphidium balchanorum (H. Kraschen.) Polj. 118 
Seriphidium baldshuanicum (H. Kraschen. & 

Zaprj.) Polj. 118 
Seriphidium barrelieri (Besser) Sojak 118 
Seriphidium bicolor (Rech. f. & Wagenitz) Bremer 

& Humphries 118 
Seriphidium bigelowii (A. Gray) Bremer & 

Humphries 118 
Seriphidium borotalense (Polj.) Ling & Y. R. Ling 

118 



Seriphidium botschantzevii (Filat.) Y. R. Ling 118 
Seriphidium brevifolium (Wallich ex DC.) Ling & 

Y. R. Ling 118 
Seriphidium caerulescens (L.) Sojak 118 
Seriphidium camelorum (H. Kraschen.) Polj. 118 
Seriphidium canum (Pursh) W. A. Weber 118 
Seriphidium chitralense (Podl.) Bremer & 

Humphries 118 
Seriphidium ciniforme (H. Kraschen. & Popov, ex 

Polj.) Polj. 118 
Seriphidium cinum (P. Bergius ex Polj.) Polj. 118 
Seriphidium compactum (Fischer ex DC.) Polj. 118 
Seriphidium cretaceum (Fiori) Bremer & 

Humphries 118 
Seriphidium densifolium (Filat.) Y. R. Ling. 118 
Seriphidium deserti (H. Kraschen.) Polj. 118 
Seriphidium diffusum (H. Kraschen. ex Polj.) Y. 

R. Ling 118 
Seriphidium dubyanskyanum (H. Kraschen. ex 

Polj.) Polj. 118 
Seriphidium dumosum (Polj.) Polj. 118 
Seriphidium dzevanovskyi (Leonova) Sojak 118 
Seriphidium elongatum (Filat. & Ladyg.) Bremer 

& Humphries in Y. R. Ling 
118 
Seriphidium eremophilum (H. Kraschen. & 

Butkov ex Polj.) Bremer & Humphries in Y. R. 

Ling 118 
Seriphidium federovii (Rzazade) Y. R. Ling 118 
Seriphidium fedtschenkoanum (H. Kraschen.) Polj. 

118 
Seriphidium ferganense (H. Kraschen. ex Polj.) 

Polj. 118 
Seriphidium finitum (Kitagawa) Ling & Y. R. Ling 

118 
Seriphidium fragrans (Willd.) Polj. 118 
Seriphidium freitagii (Podl.) Bremer & Humphries 

118 
Seriphidium fulvellum (Filat. & Ladyg.) Bremer & 

Humphries 118 
Seriphidium ghazniense (Podl.) Bremer & 

Humphries 118 
Seriphidium ghoratense (Podl.) Bremer & 

Humphries 118 
Seriphidium glanduligerum (H. Kraschen. ex Polj.) 

Polj. 118 
Seriphidium glaucinum (H. Kraschen. ex Polj.) 

Bremer & Humphries in Y. R. Ling 
118 
Seriphidium gorjaevii (Polj.) Y. R. Ling & 

Humphries 118 
Seriphidium gracilescens (H. Kraschen. & Iljin) 

Polj. 118 
Seriphidium grenardii (Franchet) Y. R. Ling 118 
Seriphidium gurganicum (H. Kraschen.) Bremer & 

Humphries in Y. R. Ling 
118 
Seriphidium gypsaceum (H. Kraschen., Popov & 

Lincz. ex Polj.) Polj. 118 
Seriphidium halophilum (H. Kraschen.) Polj. 119 
Seriphidium heptapotamicum (Polj.) Ling & Y. R. 

Ling 119 
Seriphidium herba-album (Asso) Sojak 119 
Seriphidium incultum (Del.) Y. R. Ling 119 
Seriphidium issykkulense (Polj.) Polj. 119 
Seriphidium junceum (Karelin & Kir.) Polj. 119 
Seriphidium kandaharense (Podl.) Bremer & 

Humphries 119 
Seriphidium karatavicum (H. Kraschen. & Abolin 

ex Polj.) Ling & Y, R. Ling 119 
Seriphidium kasakorum (H. Kraschen.) Bremer & 

Humphries 119 
Seriphidium kaschgaricum (H. Kraschen.) Polj. 

119 
Seriphidium kemrudicum (H. Kraschen.) Polj. 119 
Seriphidium kermanense (Podl.) Bremer & 

Humphries 119 



Seriphidium khorassanicum (Podl.) Bremer & 

Humphries 119 
Seriphidium knorringianum (H. Kraschen.) Polj. 

119 
Seriphidium kochiiforme (H. Kraschen. & Lincz. 

ex Polj.) Polj. 119 
Seriphidium kopetdaghensc (H. Kraschen. ex 

Polj.) Polj. 119 
Seriphidium korovinii (Polj.) Polj. 119 
Seriphidium korshinskyi (H. Kraschen. ex Polj.) 

Y. R. Ling 119 
Seriphidium kurramense (Oaz.) Y. R. Ling 119 
Seriphidium lehmannianum (Bunge) Polj. 119 
Seriphidium lerchianum (G. Weber ex Stechm.) 

Polj. 119 
Seriphidium lessingianum (Besser) Polj. 119 
Seriphidium leucodes (Schrenk) Polj. 119 
Seriphidium leucotrichum (H. Kraschen. ex Polj.) 

Bremer & Humphries in Y. R. Ling 119 
Seriphidium lobulifolium (Boiss.) Polj. 119 
Seriphidium longilobum (Osterh.) Bremer & 

Humphries 119 
Seriphidium maritimum (L.) Polj. 119 
Seriphidium mendozanum (DC.) Bremer & 

Humphries 119 
Seriphidium minchunense Ling & Y. R. Ling 119 
Seriphidium mogoltavicum (Polj.) Y. R. Ling 119 
Seriphidium mongolorum (H. Kraschen.) Ling & 

Y. R. Ling 119 
Seriphidium monogynum (Waldst. & Kit.) Polj. 

119 
Seriphidium mucronulatum (Polj.) Y. R. Ling 119 
Seriphidium namanganicum (Polj.) Polj. 119 
Seriphidium nigricans (Filat. & Ladyg.) Bremer & 

Humphries in Y. R. Ling 119 
Seriphidium nitrosum (G. Weber in Stechm.) Polj. 

119 
Seriphidium novum (Nelson) W. A. Weber 119 
Seriphidium nutans (Willd.) Sojak 119 
Seriphidium oliverianum (Gay ex Besser) Bremer 

& Humphries in Y. R. Ling 
119 
Seriphidium oranense (Deb. ex Filat.) Y. R. Ling 

119 
Seriphidium oratense (Deb. & Filat.) Y. R. Ling 

119 
Seriphidium palmeri (A. Gray) Bremer & 

Humphries 118, 119 
Seriphidium pauciflorum (G. Weber in Stechm.) 

Polj. 119 
Seriphidium poljakovii (Filat.) Y. R. Ling 119 
Seriphidium polystichum (Polj.) Y. R. Ling. 119 
Seriphidium porrectum (H. Kraschen. ex Polj.) 

Polj. 119 
Seriphidium prasinum (H. Kraschen. ex Polj.) 

Polj. 119 
Seriphidium prolixum (H. Kraschen. ex Polj.) Polj. 

119 
Seriphidium pygmaeum (A. Gray) W. A. Weber 

119 
Seriphidium quettense (Podl.) Bremer & 

Humphries 119 
Seriphidium rhodanthum (Rupr.) Polj. 119 
Seriphidium rigidum (Nutt.) W. A. Weber 119 
Seriphidium rothrockii (A. Gray) W. A. Weber 

119 
Seriphidium saharum (Pomel) Y. R. Ling 119 
Seriphidium saissanicum (H. Kraschen.) Bremer & 

Humphries in Y. R. Ling 
119 

Seriphidium santolinum (Schrenk) Polj. 119 
Seriphidium santonicum (L.) Sojak 119 
Seriphidium sawanense Y. R. Ling & Humphries 

119 
Seriphidium schrenkianum (Ledeb.) Polj. 119 
Seriphidium scopiforme (Ledeb.) Polj. 119 
Seriphidium scotinum (Ncvski) Polj. 119 
Seriphidium semiaridum (H. Kraschen. & 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



175 



Lavrenko) Ling & Y. R. Ling 119 
Seriphidium serotinum (Bunge) Polj. 119 
Seriphidium siebcri (Besser) Bremer & Humphries 

in Y. R. Ling 119 
Seriphidium skorniakowii (Winkler) Bremer & 

Humphries in Y. R, Ling 119 
Seriphidium spicigerum (Koch) Polj. 119 
Seriphidium stenocephalum (H. Kraschen. ex 

Polj.) Polj. 119 
Seriphidium subchrysolepis (Filat.) Bremer & 

Humphries 119 
Seriphidium sublcssingianum (Kell.) Polj. 119 
Seriphidium subsalsum (Filat.) Bremer & 

Humphries in Y. R. Ling 119 
Seriphidium szowitzianum (Besser) Polj. 119 
Seriphidium tauricum (Willd.) Polj. 119 
Seriphidium tecti-mundi (Podl.) Bremer & 

Humphries 119 
Seriphidium tenuisectum (Nevski) Polj. 119 
Seriphidium terrae-albae (H. Kraschen.) Polj. 119 
Seriphidium thomsonianum (C. B. Clarke) Ling & 

Y. R. Ling 119 
Seriphidium tianshanicum (H. Kraschen.) Y. R. 

Ling 119 
Seriphidium transiliense (Polj.) Polj. 119 
Seriphidium tridentatum (Nutt.) W. A. Weber 118, 

119 
Seriphidium tripartitum (Rydb.) W. A. Weber 120 
Seriphidium turanicum (H. Kraschen.) Polj. 120 
Seriphidium turcomanicum (Gand.) Polj. 120 
Seriphidium vachanicum (H. Kraschen. ex Polj.) 

Polj. 120 
Seriphidium validum (H. Kraschen. ex Polj.) Polj. 

120 
Seriphidium valesianum (Lam.) Y. R. Ling. 120 
Seriphidium vallesiacum (All.) Sojak 120 
Seriphidium vaseyanum (Rydb.) W. A. Weber 120 
Solenogyne Cass. 161 
Soliva Ruiz Lopez & Pavon 76, 81, 147, 150, 151, 

158, 159, 160 
Soliva anthemifolia R. Br. 160 
Soliva macrocephala Cabrera 160 
Soliva mutisii Kunth 160 
Soliva neglecta Cabrera 160 
Soliva pterosperma (Juss.) Less. 160 
Soliva sessilis Ruiz Lopez & Pavon 159, 160 
Soliva stolonifera (Brot.) R. Br. ex G. Don in 

Loudon 159, 160 
Soliva triniifolia Griseb. 160 
Soliva valdiviana Phihppi 160 
Spathipappus Tz\e\e\ 100, 101, 105 
Spathipappus chitralensis Podl. 101, 102 
Spathipappus griffithii (C. B. Clarke) Tzvelev 101, 

102 
Spathipappus porphyrostephanus (Rech. f.) 

Tzvelev 101, 103 
Sphaeroclinium (DC.) Schultz-Bip. 157, 158 
Sphaerodinium nigellifolium (DC.) Schultz-Bip. 

158 
Sphaeromeria Nutt. 111-113, 117 
Sphaeromeria argentea Nutt. 117 
Sphaeromeria cana (D. C. Eaton) A. A. Heller 

117 
Sphaeromeria capitata Nutt. 117 
Sphaeromeria compacta (H. M. Hall) A. 

Holmgren, Shultz & Lowrey 117 
Sphaeromeria diversifoha (D. C. Eaton) Rydb. 117 
Sphaeromeria martirensis (Wiggins) A. Holmgren, 

Shultz & Lowrey 117 
Sphaeromeria potentilloides (A. Gray) A. A. 

Heller 117 
Sphaeromeria ruthiae A. Holmgren. Shultz & 

Lowrey 117 
Sphaeromeria simplex (Nelson) A. A. Heller 117 
Sphenogyne R. Br. 93, 
Sphenogyne brachyloba Kunze 93, 
Stenops B. Nord. 161 
Stilpnolepis H. Kraschen. Ill, 112, 116 



Stilpnolepis centiflora (Maxim.) H. Kraschen. 116 
Stilpnolepis intricata (Franchet) Shih 116 
Stilpnophylon Less. 95 
Stilpnophyton inopinatum Hutch. 96 
Stilpnophylon linifolium (L. f.) Less. 96 
Stilpnophyton longifolium (Thunb.) Less. 96 
Stilpnophyton oocephalum DC. 96 



Tanacetinae Bremer & Humphries 

73, 74, 76, 77, 84, 96, 99, 100, 107, 138, 145 

Tanacetopsis (Tzvelev) Kovalevsk. 77, 99, 104, 

106, 109 
Tanacetopsis afghanica (Gilli) Bremer & 

Humphries 104 
Tanacetopsis botschantzevii (Kovalevsk.) 

Kovalevsk. 104 
Tanacetopsis doabensis (Podl.) Bremer & 

Humphries 104 
Tanacetopsis eriobasis (Rech. f.) Kovalevsk. 104 
Tanacetopsis ferganensis (Kovalevsk.) Kovalevsk. 

104 
Tanacetopsis freitagii (Podl.) Bremer & Humphries 

104 
Tanacetopsis golovskovii (Polj.) Karmysch. 104 
Tanacetopsis hedge! (Podl.) Bremer & Humphries 

104 
Tanacetopsis kamelinii Kovalevsk. 104 
Tanacetopsis karataviensis (Kovalevsk.) 

Kovalevsk. 104 
Tanacetopsis kjurendaghii Kurbanov 104 
Tanacetopsis krascheninnikovii (Nevski) 

Kovalevsk. 104 
Tanacetopsis mucronata (Regel & Schmalh.) 

Kovalevsk. 104 
Tanacetopsis pjataeviae (Kovalevsk.) Karmysch. 

104 
Tanacetopsis platyrachis (Boiss.) Kovalevsk. 104 
Tanacetopsis santoana (H. Kraschen., Popov & 

Vved.) Kovalevsk. 104 
Tanacetopsis setacea (Regel & Schmalh.) 

Kovalevsk. 104 
Tanacetopsis submarginata (Kovalevsk.) 

Kovalevsk. 104 
Tanacetopsis subsimilis (Rech. f.) Kovalevsk. 104 
Tanacetopsis tripinnatifida (Oliver) Kovalevsk. 105 
Tanacetopsis urgutensis (Popov) Kovalevsk. 105 
Tanacetum L. 
76, 77, 96, 97, 98, 99, 100-104, 105-111, 113, 114, 

117, 128, 134, 140 
Tanacetum sect. Asterotricha Tzvelev 101 
Tanacetum sect. Balsamita (Miller) Schultz-Bip. 

101 
Tanacetum sect. Hippolytia (Polj.) Podl. 106 
Tanacetum sect. Pyrethrum (Zinn.) Reichenb.f. 

98, 140 
Tanacetum sect. Pyrethrum subsect. 

Leucanthemopsis Giroux 101 
Tanacetum abrotanifolium (L.) Druce 101 
Tanacetum abrotanoides Bremer & Humphries 101 
Tanacetum achilleifolium (M. Bieb.) Schultz-Bip. 

101 
Tanacetum afghanicum (Gilli) Podl. 104 
Tanacetum akinfievii (Alexej.) Tzvelev 101 
Tanacetum alashanense Ling 106 
Tanacetum alatavicum Herder 101 
Tanacetum albipannosum Huber-Mor. & Grierson 

101 
Tanacetum alyssifolium (Bornm.) Grierson 101 
Tanacetum annuum L. 100, 101 
Tanacetum arctodzhungaricum (Golosk.) Bremer 

& Humphries 101 
Tanacetum archibaldii Podl. 101 
Tanacetum argenteum (Lam.) Willd. 101 
Tanacetum argyranthemoides (Boiss. & Kotschy) 

Schultz-Bip. 101 
Tanacetum argyrophyllum (C. Koch) Tzvelev 103 
Tanacetum armenum (DC.) Schultz-Bip. 101 



Tanacetum aromaticum (Tzvelev) Bremer & 

Humphries 101 
Tanacetum artemisioides Schultz-Bip. in Hook. f. 

101 
Tanacetum atkinsonii (C. B. Clarke) Kitam. 101 
Tanacetum aucheranum (DC.) Schultz-Bip. 101 
Tanacetum aucheri DC. 102 
Tanacetum audibertii (Req.) DC. 102 
Tanacetum balsamita L. 101, 102 
Tanacetum bamianicum Podl. 102 
Tanacetum barclayanum DC. 102 
Tanacetum bipinnatum (L.) Schultz-Bip. 102 
Tanacetum boreale Fischer ex DC. 102 
Tanacetum budjurnense (Rech. f.) Tzvelev 102 
Tanacetum cadmeum (Boiss.) Heyw. 102 
Tanacetum camphoratum Less. 101, 102 
Tanacetum canescens DC. 102 
Tanacetum cappadocicum (DC.) Schultz-Bip. 102 
Tanacetum changaicum (H. Kraschen. ex Grubov) 

Bremer & Humphries 102 
Tanacetum chiliophyllum (Fischer & C. Meyer) 

Schultz-Bip. 102 
Tanacetum chitralense (Podl.) Bremer & 

Humphries 102 
Tanacetum cilicium (Boiss.) Grierson 102 
Tanacetum cinerariifolium (Trevir) Schultz-Bip. 

101, 102 
Tanacetum coccineum (Willd.) Grierson 101, 102 
Tanacetum compactum Hall 117 
Tanacetum corymbiforme (Tzvelev) Bremer & 

Humphries 102 
Tanacetum corymbosum (L.) Schultz-Bip. 100, 

101, 102 
Tanacetum crassicollum (Rech. f.) Podl. 106 
Tanacetum crassipes (Stchegl.) Tzvelev 102 
Tanacetum czerniakowskae (H. Kraschen.) Parsa 

102 
Tanacetum daghestanicum (Rupr. ex Boiss.) 

Bremer & Humphries 102 
Tanacetum demetrii (Manden.) Bremer & 

Humphries 102 
Tanacetum densum (Labill.) Schultz-Bip. 102 
Tanacetum depauperatum (Post) Grierson 101, 102 
Tanacetum djilgense (Franchet) Podl. 98 
Tanacetum doabense Podl. 104 
Tanacetum dolomiticum (Galushko) Bremer & 

Humphries 102 
Tanacetum dolichophyllum (Kitam.) Kitam. 106 
Tanacetum duderanum (Boiss.) Tzvelev 103 
Tanacetum dumosum Boiss. 102 
Tanacetum eginense (Hausskn. ex Bornm.) 

Grierson 102 
Tanacetum elongatum (Bornm. & Gauba) Parsa 

102 
Tanacetum eriobasis (Rech. f.) Kovalevsk. 104 
Tanacetum falcatolobatum H. Kraschen. 102 
Tanacetum falconcri Hook. f. 102 
Tanacetum ferulaceum (Webb ex Berth.) 

Schultz-Bip. 102, 107 
Tanacetum flavovirens (Boiss.) Tzvelev 103 
Tanacetum freitagii Pod\. 104 
Tanacetum funkii Schultz-Bip. ex Willk. 102 
Tanacetum galae (Popov) Nevski 102 
Tanacetum gaiushkoi (Prima) Bremer & 

Humphries 102 
Tanacetum germanicopolitanum (Bornm. & 

Heimerl) Grierson 102 
Tanacetum ghoratense Podl. 102 
Tanacetum gilletii Podl. 105 
Tanacetum gilliatii (Turrill) Parsa 102 
Tanacetum glabrum (Decne) Muradyan 98 
Tanacetum glanduliferum (Sommier & Levier) 

Bremer & Humphries 102 
Tanacetum griffithii (C. B. Clarke) Muradyan 102 
Tanacetum grossheimii (Sosn.) Muradyan 102 
Tanacetum haradjanii (Rech. f.) Grierson 101, 102 
Tanacetum haussknechtii (Bornm.) Grierson 102 
Tanacetum hedgei Podl. 104 



176 



K. BREMER AND C. J. HUMPHRIES 



Tanacetum heimerli (Nab.) Parsa 102 
Tanacetum herderi Kegel & Schmal. 102 
Tanacetum heterophyllum Boiss. 103 
Tanacetum heterotomum (Bornm.) Grierson 102 
Tanacetum himachalense Aswal & Mehrotra 106 
Tanacetum hissaricum (H. Kraschen.) Bremer & 

Humphries 102 
Tanacetum hololeucum (Bornm.) Podl. 102 
Tanacetum huronense Nutt. 102 
Tanacetum junesarense (Bornm.) Parsa 103 
Tanacetum karelinianum Bremer & Humphries 102 
Tanacetum karelinii Tzvelev 102 
Tanacetum kaschgarianum Bremer & Humphries 

102 
Tanacetum kaschgaricum H. Kraschen. 102, 106 
Tanacetum kelleri (Krylov & Plotn.) Takht. 102 
Tanacetum khorassanicum (H. Kraschen.) Parsa 

102 
Tanacetum kittaryanum (C. Meyer) Tzvelev 103 
Tanacetum komarovii (Winkler) Muradyan 103, 

105 
Tanacetum kotschyi (Boiss.) Grierson 102 
Tanacetum krylovianum (H. Kraschen.) Bremer & 

Humphries 102 
Tanacetum kubense (Grossh.) Muradyan 102 
Tanacetum lanuginosum Schultz-Bip. & Herder 

102 
Tanacetum ledebourii Sc\\\AXz-E'\\). 153, 154 
Tanacetum leptophyllum (Steven) Schultz-Bip. 

102, 104 
Tanacetum longipedunculatum (Sosn.) Tzvelev 102 
Tanacetum macrocephalum Pampan. 102 
Tanacetum macrophyllum (Waldst. & Kit.) 

Schultz-Bip. 101, 102 
Tanacetum macropodum Hemsley & Lace 105 
Tanacetum marionii (Albov) Bremer & Humphries 

102 
Tanacetum maymanense Podl. 103 
Tanacetum microphyllum DC. 101, 103 
Tanacetum mikeschinii (Tzvelev) Takht. 103 
Tanacetum millefolium (L.) Tzvelev 101, 103 
Tanacetum modestum (Heimerl ex Stapf) Parsa 

102 
Tanacetum mucroniferum Huber-Mor. & Grierson 

103 
Tanacetum mucronulatum (Hoffsgg & Link) 

Heyw. 103 
Tanacetum myriophyllum Willd. 103 
Tanacetum nitens (Boiss. & Noe) Grierson 103 
Tanacetum nivale Schultz-Bip. 103 
Tanacetum niveum (Lagasca) Schultz-Bip. 103 
Tanacetum nuristanicum Podl. 103 
Tanacetum nuratavicum (H. Kraschen.) Muradyan 

105 
Tanacetum odessanum (Klokov) Tzvelev 103 
Tanacetum oligocephalum (DC.) Schultz-Bip. 102 
Tanacetum oltense (Sosn.) Grierson 103 
Tanacetum ordubadense (Manden.) Bremer & 

Humphries 103 
Tanacetum oxylepis (Bordz.) Grierson 103 
Tanacetum oxystegium (Sosn.) Grierson 103 
Tanacetum paczoskii (Zef.) Tzvelev 103 
Tanacetum paghmanense Podl. 105 
Tanacetum pakistanicum Podl. 103 
Tanacetum paleaceum Podl. 103 
Tanacetum paradoxum Bornm. 103 
Tanacetum parthenifolium (Willd.) Schultz-Bip. 

103 
Tanacetum parthenium (L.) Schultz-Bip. 101, 103 
Tanacetum petiolosum Pampan. 103 
Tanacetum petrareum (Shih) Bremer & Humphries 

103 
Tanacetum peucedanifolium (Sosn.) Bremer & 

Humphries 103 
Tanacetum pinnatum Boiss. 103 
Tanacetum podlechii Bremer & Humphries 103 
Tanacetum polyccphalum Schultz-Bip. 103 



Tanacetum porphyrostephanum (Rech. f.) Bremer 

& Humphries 103 
Tanacetum poteriifolium (Ledeb.) Grierson 103 
Tanacetum praeteritum (Horw.) Heyw. 103 
Tanacetum pseudoachillea Winkler 103 
Tanacetum ptarmiciflorum (Webb & Berth.) 

Schultz-Bip. 103, 107 
Tanacetum pulchellum (Turcz.) Schultz-Bip. 103 
Tanacetum pulchrum (Ledeb.) Schultz-Bip. 103 
Tanacetum punctatum (Desr.) Grierson 103 
Tanacetum pyrethroides (Karelin & Kir.) 

Schultz-Bip. 98 
Tanacetum richterioides (Winkler) Bremer & 

Humphries 103 
Tanacetum robustum Hook. f. & Thomson ex C. 

B. Clarke 103 
Tanacetum roseum (Adams) Schultz-Bip. 101 , 103 
Tanacetum roylei (DC.) Podl. 103 
Tanacetum salsugineum Podl. 103 
Tanacetum sanguineum (Parsa) Parsa 103 
Tanacetum santolina Winkler 103 
Tanacetum saxicolum (H. Kraschen.) Tzvelev 103 
Tanacetum sclerophyllum (H. Kraschen.) Tzvelev 

103 
Tanacetum scopulorum (H. Kraschen.) Tzvelev 

103 
Tanacetum semenovii Herder 103 
Tanacetum sericeum (Adams) Schultz-Bip. 103 
Tanacetum sevanense (Sosn.) Bremer & 

Humphries 103 
Tanacetum silaifolium (Steven) Schultz-Bip. 103 
Tanacetum silvicola Podl. 103 
Tanacetum sinaicum (Fresen.) Del. ex Bremer & 

Humphries 103 
Tanacetum sipikorense (Bornm.) Grierson 103 
Tanacetum songaricum (Tzvelev) Bremer & 

Humphries 103 
Tanacetum sorbifolium (Boiss.) Grierson 103 
Tanacetum stapfianum (Rech. f.) Podl. 103 
Tanacetum subsimile (Rech. f.) Kovalevsk. 105 
Tanacetum tabrisianum (Boiss.) Sosn. & Takht. 

103 
Tanacetum tamrutense (Sosn.) Sosn. 103 
Tanacetum tanacetoides (DC.) Tzvelev 103 
Tanacetum tatsienense (Bureau & Franchet) 

Bremer & Humphries 103, 106 
Tanacetum tenuisectum (Boiss.) Podl. 103 
Tanacetum tenuissimum (Trautv.) Gross. 103 
Tanacetum tirinense Podl. 103 
Tanacetum tomentellum (Boiss.) Grierson 101, 103 
Tanacetum tomentosum (Decne) Muradyan 98 
Tanacetum trichophyllum (Sosn.) Bremer & 

Humphries 103 
Tanacetum trifoholatum Podl. 103 
Tanacetum turcomanicum (H. Kraschen.) Tzvelev 

103 
Tanacetum turlanicum (Pavlov) Tzvelev 102 
Tanacetum ulutavicum Tzvelev 103 
Tanacetum uniflorum (Fischer & C. Meyer ex 

DC.) Schultz-Bip. 104 
Tanacetum uralense (H. Kraschen.) Tzvelev 104 
Tanacetum vahlii DC. 104 
Tanacetum vulgare L. 100, 101, 104 
Tanacetum waited (Winkler) Tzvelev 104 
Tanacetum willkommii Schultz-Bip. 104 
Tanacetum yabrudae (Mout.) Charpin & Dittrich 

104 
Tanacetum zahlbruckneri (Nab.) Grierson 104 
Thaminophyllinae Bremer & Humphries 74, 76, 77, 

83, 128, 144, 145 
Thaminophyllum Harvey 144, 145, 146 
Thaminophyllum latifolium Bond 146 
Thaminophyllum multiflorum Harvey 146 
Thaminophyllum mundtii Harvey 146 
Trichanthemis Regel & Schmalh. 96, 97, 98, 105 
Trichanthemis afghanica Podl. 97 
Trichanthemis aulieatensis (B. Fedtsch.) H. 

Kraschen. 97 



Trichanthemis aurea H. Kraschen. 97 
Trichanthemis butkovii Kovalevsk. 97 
Trichanthemis karataviensis Regel & Schmalh. 97, 

99 
Trichanthemis litwinowii (H. Kraschen.) Tzvelev 

97 
Trichanthemis paradoxos (Winkler) Tzvelev 97 
Trichanthemis radiata H. Kraschen. & Vved. 97 
Trichanthemis simulans (Pavlov) Pavlov 97 
TridactyHna (DC.) Schultz-Bip. 110, 114, 115 
Tridactylina kirilowii (Turcz. ex DC.) Schultz-Bip. 

115 
Tripleurospermum Schultz-Bip. 76, 77, 83, 134, 

147-155, 155, 156 
Tripleurospermum ambiguum (Ledeb.) Franchet & 

Savat. 156 
Tripleurospermum aromaticum Rupr. ex Boiss. 101 
Tripleurospermum auriculatum (Boiss.) Rech. f. 

155, 156 
Tripleurospermum australe Pobed. 156 
Tripleurospermum baytopianum E. Hossain 156 
Tripleurospermum breviradiatum (Ledeb.) Pobed. 

156 
Tripleurospermum callosum (Boiss. & Heldr.) E. 

Hossain 156 
Tripleurospermum caucasicum (Willd.) Hayek 156 
Tripleurospermum colchicum (Manden) Pobed. 

156 
Tripleurospermum conoclinium (Boiss. & Balansa) 

Hayek 156 
Tripleurospermum corymbosum E. Hossain 156 
Tripleurospermum daghestanlcum (Rupr. ex 

Boiss.) Bremer & Humphries 156 
Tripleurospermum decipiens (Fischer & C. Meyer) 

Bornm. 156 
Tripleurospermum disciforme (C. Meyer) 

Schultz-Bip. 156 
Tripleurospermum elongatum (Fischer & C. Meyer 

ex DC.) Bornm. 156 
Tripleurospermum fissurale (Sosn.) E. Hossain 156 
Tripleurospermum froedinii Rech. f. 156 
Tripleurospermum grossheimii (Fed.) Pobed. 156 
Tripleurospermum heterolepis (Freyn & Sint.) 

Bornm. 156 
Tripleurospermum homogamum G. X. Fu 156 
Tripleurospermum hygrophilum (Bornm.) Bornm. 

156 
Tripleurospermum inodorum (L.) Schultz-Bip. 153, 

154, 156 
Tripleurospermum karjaginii (Manden. & Sof.) 

Pobed. 156 
Tripleurospermum kotschyi (Boiss.) E. Hossain 

156 
Tripleurospermum lesbiacum (Candargy) Rech. f. 

156 
Tripleurospermum limosum (Maxim.) Pobed. 156 
Tripleurospermum maritimum (L.) K. Koch 155, 

156 
Tripleurospermum maritimum (L.) K. Koch ssp. 

maritimum 156 
Tripleurospermum maritimum ssp. phaeocephalum 

(Rupr.) Rauschert 156 
Tripleurospermum maritimum ssp. subpolare 

(Pobed.) Rauschert 156 
Tripleurospermum microcephalum (Boiss.) 

Bornm. 156 
Tripleurospermum monticolum (Boiss. & Huet) 

Bornm. 156 
Tripleurospermum oreades (Boiss.) Rech.f. 156 
Tripleurospermum parviflorum (Willd.) Pobed. 

156 
Tripleurospermum perforatum (Merat) Lainz 155, 

156 
Tripleurospermum phaeocephalum (Rupr.) Pobed. 

156 
Tripleurospermum pichleri (Boiss.) Bornm. 156 
Tripleurospermum repens (Freyn & Sint.) Bornm. 

156 



GENERIC MONOGRAPH OF ASTERACEAE-ANTHEMIDEAE 



177 



Tripleurospcrmum rosellum (Boiss. & Orph.) Urs 

Hayek 156 Ursi 

Tripleurospcrmum rupestre (Sommier & Levier) Urs 

Pobed. 156 Urs 

Tripleurospermum sevanense (Manden.) Pobed. Urs 

156 Urs 

Tripleurospermum subnivale Pobed. 156 Ursi 

Tripleurospermum subpolare Pobed. 156 Ursi 

Tripleurospermum szowitzii (DC.) Pobed. 156 Ursi 

Tripleurospermum tchihatchewii (Boiss.) Bornm. Urs 

156 Ursi 

Tripleurospermum tempskyanum (Freyn & Sint.) Ursi 

Hayek 156 Ursi 

Tripleurospermum tcnuifolium (Kit.) Freyn. 156 Ursi 

Tripleurospermum tctragonospermum (Schum.) Ursi 

Pobed. 156 Ursi 

Tripleurospermum transcaucasicum (Manden.) Ursi 

Pobed. 156 Ursi 

Tripleurospermum tridentatum Hoffm. 143 Ursi 

Tripleurospermum tzvelcvii Pobed. 156 Urs 

Turaniphytum Polj. 113, 121, 125 Ursi 

Turaniphytum codringtonii (Rech. f.) Podl. 125 Urs: 

Turaniphytum cranthcmum (Bunge) Polj. 125 Urs; 

Turaniphytum kopetdaghense Po\\. 125 Ursi 

Urs: 

Ugamia Pavlov 96, 97, 98 Urs 

Ugamia angrenica (H. Kraschen.) Tzvclev 98 Urs: 

Ugamia trichanthemoides Pavlov 98 Urs 

Ursinia Gaertn. 75, 77, 83, 91, 93 Urs: 

Ursinia abrotanifolia (R. Br.) Sprengel 93 Urs 

Ursinia anethoides (DC.) N. E. Br. 93 Urs 

Ursinia anthemoides (L.) Poiret 93 Urs 

Ursinia brachyloba (Kunze) Bremer & Humphries Urs 

93 Urs 

Ursinia cakilefolia DC. 93 Urs, 
Ursinia caledonica (E. Phillips) Prassler 93 



nia calcnduliflora (DC.) N. E. Br. 93 

nia chrysanthemoides (Less.) Harvey 91 

nia coronopifolia (Less.) N. E. Br. 93 

nia crithmoides (P. Bergius) Poiret 93 

nia dentata (L.) Poiret 93 

nia discolor (DC.) N. E. Br. 93 

nia drcgeana (DC.) N. E. Br. 93 

nia eckloniana (Sondcr) N. E. Br. 93 

nia filipes (E. Meyer ex DC.) N. E. Br. 93 

nia frutcscens Dinter 93 

nia hcterodonta (DC.) N. E. Br. 93 

nia hispida (DC.) N. E. Br. 93 

nia macropoda (DC.) N. E. Br. 93 

nia mcrxmucllcri Prassler 93 

nia montana DC. 93 

nia nana DC. 93 

nia nudicaulis (Thunb.) N. E. Br. 93 

nia orcogena Schltr ex Prassler 93 

nia paleacca (L.) Mocnch 93 

nia paradoxa Gaertn. 91 , 93 

nia pilifera (P. Bergius) Poiret 93 

nia pinnata (Thunb.) Prassler 93 

nia punctata (Thunb.) N. E. Br. 93 

nia pygmaea DC. 93 

nia quinquepartita (DC.) N. E. Br. 93 

nia rigidula (DC.) N. E. Br. 93 

nia saxatilis N. E. Br. 93 

nia scariosa (Alton) Poiret 93 

nia sericea (Thunb.) N. E. Br. 93 

nia serrata (L. f.) Poiret 93 

nia speciosa DC. 93 

nia subflosculosa (DC.) Prassler 93 

nia tenuifolia (L.) Poiret 93 

nia trifida (Thunb.) N. E. Br. 93 

nieae H. Robinson & Brettell 75 



Ursiniinae Bremer & Humphries 75-77, 80, 81, 83, 
93 91-93 

Ursiniopsis E. Phillips 93, 

Vesicarpa Rydb. 1 17 

Vesicarpa polentilloides (A. Gray) Rydb. 117 



Waldheimia Karelin & Kir. 98 
Waldheimia glabra (Decnc) Regel 98 
Waldheimia huegelii (Schultz-Bip.) Tzvelev 98 
Waldheimia lasiocarpa G. X. Fu 98 
Waldheimia nivea (C. B. Clarke) Rcgel 98 
Waldheimia stoliczkae (C. B. Clarke) Ostenf. 98 
Waldheimia tomentosa (Dccne) Rcgel 98 
Waldheimia transalaica Tzvelev 98 
Waldheimia tridactylites Karelin & Kir. 98 
Waldheimia veslila (C. B. Clarke) Pampan. 98 

Xylanthemum Tzvelev 1(X), 101, 105 
Xylanthemum fisherae (Aitch. & Hemsley) 

Tzvclev 105 
Xylanthemum gilletii (Podl.) Bremer & Humphries 

105 
Xylanthemum lingulatum (Boiss.) Bremer & 

Humphries 105 
Xylanthemum macropodum (Hemsley & Lace) 

Bremer & Humphries 105 
Xylanthemum paghmanense (Podl.) Bremer & 

Humphries 105 
Xylanthemum pamiricum (Hoffm.) Tzvelev 105 
Xylanthemum polycladum (Rech. f.) Tzvelev 105 
Xylanthemum rupestre (Popov ex Nevski) Tzvelev 

105 
Xylanthemum tianshanicum (H. Kraschen.) 

Muradyan97, 100, 105, 111 



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Woelkerling and S.J. Campbell. 1992. Pp. 1-107. 63 
figs. £37.00 

No. 2 Palynological evidence for the generic delimitation of 

Sechium (Cucurbitaceae) and its allies. J.L. Alvarado. R. 

Lira-Saade & J. Caballero. 1992. Pp. 109-121. 

Seaweeds of the western coast of tropical Africa and 

adjacent islands: a critical assessment. IV. Rhodophyta 

(Florideae) 3. Genera H-K. J.H. Price. D.M. John & 

G.W. Lawson. 1992. pp. 123-146. 

Two new species of Solarium section Geminata 

(Solanaceae) from Cerro del Torra in western Colombia. 

S. Knapp. 1992. Pp. 147-152. 

Fissidens ceylonensis Dozy & Molkenb. (Musci: 

Fissidentaceae) and some allied taxa from southern 

India. L.T. EUis. 1992. Pp. 153-156, 2 figs. 

New species of Piper (Piperaceae) from central America. 

M. Tebbs. 1992. Pp. 157-158. 

Studies on the Cretan flora 1. Floristic notes. N.J. 

Turland. 1992. Pp. 159-164. 

Studies on the Cretan flora 2. The Dianthus juniperintis 

complex (Caryophyllaceae). N.J. Turland. 1992. 

Pp. 165-169. £37.50 



COJiTgJITS 

55 New taxa of Gentiana (Gentianaceae) from Western China and the Himalayan region 

T.-N. Ho and S.-W. Uu 
61 New combinations, names and taxonomic notes on Gentianella (Gentianaceae) from South 

America and New Zealand 

T.-N. Ho and S.-W. Uu 
67 Studies in Hypericum: validation of new names 

N.K.B. Robson 
71 Generic monograph of the Asteraceae-Anthemideae 

K. Bremer and C.J. Humphries 



Bulletin of The Natural yigtpf^,, 

BOTANY SERIES 

Vol. 23, No. 2, November 1993